forked from pensoft/ZooKeys-xml
/
10.3897_zookeys.100.1537.xml
1262 lines (1262 loc) · 134 KB
/
10.3897_zookeys.100.1537.xml
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
209
210
211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
252
253
254
255
256
257
258
259
260
261
262
263
264
265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
302
303
304
305
306
307
308
309
310
311
312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
355
356
357
358
359
360
361
362
363
364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
410
411
412
413
414
415
416
417
418
419
420
421
422
423
424
425
426
427
428
429
430
431
432
433
434
435
436
437
438
439
440
441
442
443
444
445
446
447
448
449
450
451
452
453
454
455
456
457
458
459
460
461
462
463
464
465
466
467
468
469
470
471
472
473
474
475
476
477
478
479
480
481
482
483
484
485
486
487
488
489
490
491
492
493
494
495
496
497
498
499
500
501
502
503
504
505
506
507
508
509
510
511
512
513
514
515
516
517
518
519
520
521
522
523
524
525
526
527
528
529
530
531
532
533
534
535
536
537
538
539
540
541
542
543
544
545
546
547
548
549
550
551
552
553
554
555
556
557
558
559
560
561
562
563
564
565
566
567
568
569
570
571
572
573
574
575
576
577
578
579
580
581
582
583
584
585
586
587
588
589
590
591
592
593
594
595
596
597
598
599
600
601
602
603
604
605
606
607
608
609
610
611
612
613
614
615
616
617
618
619
620
621
622
623
624
625
626
627
628
629
630
631
632
633
634
635
636
637
638
639
640
641
642
643
644
645
646
647
648
649
650
651
652
653
654
655
656
657
658
659
660
661
662
663
664
665
666
667
668
669
670
671
672
673
674
675
676
677
678
679
680
681
682
683
684
685
686
687
688
689
690
691
692
693
694
695
696
697
698
699
700
701
702
703
704
705
706
707
708
709
710
711
712
713
714
715
716
717
718
719
720
721
722
723
724
725
726
727
728
729
730
731
732
733
734
735
736
737
738
739
740
741
742
743
744
745
746
747
748
749
750
751
752
753
754
755
756
757
758
759
760
761
762
763
764
765
766
767
768
769
770
771
772
773
774
775
776
777
778
779
780
781
782
783
784
785
786
787
788
789
790
791
792
793
794
795
796
797
798
799
800
801
802
803
804
805
806
807
808
809
810
811
812
813
814
815
816
817
818
819
820
821
822
823
824
825
826
827
828
829
830
831
832
833
834
835
836
837
838
839
840
841
842
843
844
845
846
847
848
849
850
851
852
853
854
855
856
857
858
859
860
861
862
863
864
865
866
867
868
869
870
871
872
873
874
875
876
877
878
879
880
881
882
883
884
885
886
887
888
889
890
891
892
893
894
895
896
897
898
899
900
901
902
903
904
905
906
907
908
909
910
911
912
913
914
915
916
917
918
919
920
921
922
923
924
925
926
927
928
929
930
931
932
933
934
935
936
937
938
939
940
941
942
943
944
945
946
947
948
949
950
951
952
953
954
955
956
957
958
959
960
961
962
963
964
965
966
967
968
969
970
971
972
973
974
975
976
977
978
979
980
981
982
983
984
985
986
987
988
989
990
991
992
993
994
995
996
997
998
999
1000
<!DOCTYPE article PUBLIC "-//TaxonX//DTD Taxonomic Treatment Publishing DTD v0 20100105//EN" "tax-treatment-NS0.dtd"><article article-type="research-article" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xmlns:tp="http://www.plazi.org/taxpub">
<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">ZooKeys</journal-id>
<journal-title-group>
<journal-title>ZooKeys</journal-title>
</journal-title-group>
<issn pub-type="ppub">1313-2989</issn>
<issn pub-type="epub">1313-2970</issn>
<publisher>
<publisher-name>Pensoft Publishers</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3897/zookeys.100.1537</article-id>
<title-group>
<article-title>Changes in salt-marsh carabid assemblages after an invasion by the native grass <italic>Elymus athericus</italic> (Link) Kerguélen</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" xlink:type="simple">
<name name-style="western">
<surname>Georges</surname>
<given-names>Anita</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref></contrib>
<contrib contrib-type="author" xlink:type="simple">
<name name-style="western">
<surname>Fouillet</surname>
<given-names>Philippe</given-names>
</name>
<xref ref-type="aff" rid="A2">2</xref></contrib>
<contrib contrib-type="author" xlink:type="simple">
<name name-style="western">
<surname>Pétillon</surname>
<given-names>Julien</given-names>
</name>
<xref ref-type="aff" rid="A2">2</xref>
<xref ref-type="aff" rid="A3">3</xref>
</contrib>
</contrib-group>
<aff id="A1">
<label>1</label>U.M.R. 6553 – ‘‘Ecosystèmes, Biodiversité, Evolution’’, Université de Rennes 1, 263 Avenue du Gal Leclerc, CS74205, 35042 Rennes Cedex, France
</aff>
<aff id="A2">
<label>2</label>U.R.U. 420 – ‘‘Biodiversité fonctionnelle et Gestion des territoires’’, Université de Rennes 1, 263 Avenue du Général Leclerc, CS 74205, 35042 Rennes Cedex, France
</aff>
<aff id="A3">
<label>3</label>Evolutionary Ecology Group, University of Antwerp, Groenenborgerlaan 171, 2020 Antwerpen, Belgium
</aff>
<author-notes>
<fn fn-type="corresp">
<p>Corresponding author: Julien Pétillon (<email xlink:type="simple">julien.petillon@univ-rennes1.fr</email>).</p>
</fn>
<fn fn-type="edited-by">
<p>Academic editor: J. Noordijk</p>
</fn>
</author-notes>
<pub-date pub-type="collection">
<year>2011</year>
</pub-date>
<pub-date pub-type="epub">
<day>20</day>
<month>5</month>
<year>2011</year>
</pub-date>
<issue>100</issue>
<fpage>407</fpage>
<lpage>419</lpage>
<history>
<date date-type="received">
<day>01</day>
<month>12</month>
<year>2009</year>
</date>
<date date-type="accepted">
<day>01</day>
<month>4</month>
<year>2010</year>
</date>
</history>
<permissions>
<copyright-statement>Anita Georges, Philippe Fouillet, Julien Pétillon</copyright-statement>
<license license-type="creative-commons-attribution" xlink:href="http://creativecommons.org/licenses/by/3.0" xlink:type="simple">
<license-p>This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p>
</license>
</permissions>
<abstract>
<label>Abstract</label>
<p>As a result of an invasion by the native grass <italic><tp:taxon-name>Elymus athericus</tp:taxon-name></italic> (Link) Kerguélen (Poaceae) in the last 10 years, a major change in vegetation cover has occurred in salt marshes of the Mont Saint-Michel bay, Western France. The impact of such an invasion on carabid assemblages, a dominant group of terrestrial arthropods in these habitats and containing several stenotopic species, is investigated here. In our study site, carabid data are available from 1983 and 1984, allowing a comparison of species distribution ranges in salt marshes before (1983–1984) and after (2002) the <italic><tp:taxon-name>Elymus athericus</tp:taxon-name></italic> invasion. A total of 16,867 adults belonging to 40 species were caught. By considering the presence-absence of species shared between studies, we show that the invasion by <italic><tp:taxon-name>Elymus athericus</tp:taxon-name></italic> promoted the progression of non-coastal species (mainly <italic><tp:taxon-name>Pterostichus</tp:taxon-name></italic> s.l. spp.). This did however not interfere with resident species distributions, finally resulting in higher carabid species richness in the entire area. The species composition and abundances of carabid assemblages were also compared between natural and invaded stations in 2002. The main result is that abundances of some halophilic species decreased in one invaded plot (in case of <italic><tp:taxon-name>Pogonus chalceus</tp:taxon-name></italic> (Marsham 1802)) whereas the opposite pattern was observed for other species (e.g., <italic><tp:taxon-name>Bembidion minimum</tp:taxon-name></italic> (Fabricius 1792)). Invaded habitats were characterized by lower percentages of halophilic species and higher total species richness.</p>
</abstract>
<kwd-group>
<label>Keywords</label>
<kwd>Coleoptera</kwd>
<kwd>Carabidae</kwd>
<kwd>native invasive species</kwd>
<kwd>salt marsh</kwd>
<kwd>ecological indicators</kwd>
</kwd-group>
</article-meta>
</front>
<body>
<sec sec-type="Introduction"><title>Introduction</title>
<p>Intertidal salt marshes are ecosystems located between land and sea, undergoing periodical flooding during tides, occurring around twice a month in West-Europe. This creates some special habitat conditions, and marsh plants and animals often have special adaptations to cope with these. Salt-marsh arthropods are able to withstand floods and salinity by physiological, behavioural or morphological adaptations (e.g., <xref ref-type="bibr" rid="B13">Foster and Treherne 1976</xref>, <xref ref-type="bibr" rid="B19">Irmler et al. 2002</xref>, <xref ref-type="bibr" rid="B33">Pétillon et al. 2009</xref>). Salt marshes are among the rarest habitats in the world, covering less than 0.01% of the Earth’s surface (<xref ref-type="bibr" rid="B8">Desender and Maelfait 1999</xref>, <xref ref-type="bibr" rid="B22">Lefeuvre et al. 2003</xref>). In Europe, their surface strongly declined during the last decades, reinforcing the conservation interest in their original flora and fauna (<xref ref-type="bibr" rid="B2">Bakker et al. 2002</xref>). There is thus an urgent need to study human impacts that can either threaten (by e.g., over-grazing or habitat destruction), or enhance (by appropriate management) halophilic species in salt marshes (<xref ref-type="bibr" rid="B15">Goeldner-Gianella 1999</xref>, <xref ref-type="bibr" rid="B1">Adam 2002</xref>).</p>
<p>More recently, salt marshes have been invaded in many West-European sites by the nitrophilous grass <italic><tp:taxon-name>Elymus athericus</tp:taxon-name></italic> (<tp:taxon-name>Poaceae</tp:taxon-name>) (<xref ref-type="bibr" rid="B47">Valéry et al. 2004</xref>), probably due to increases in soil nitrogen (via the accumulation of nitrogenous compounds in the plant: <xref ref-type="bibr" rid="B23">Leport et al. 2006</xref>) and/or to the abandonment of agricultural practises (e.g., <xref ref-type="bibr" rid="B11">Esselink et al. 2000</xref>). Although <italic><tp:taxon-name>Elymus athericus</tp:taxon-name></italic> is a native species in Europe (<xref ref-type="bibr" rid="B3">Bockelmann and Neuhaus 1999</xref>) – usually growing in the upper parts of salt marshes – it can form dense, mono-specific stands, which corresponds to an invasion. This is likely to modify biodiversity and consequently ecosystem proprieties and functions as well as the conservation value of invaded areas (<xref ref-type="bibr" rid="B48">Valéry et al. 2009</xref>). Invaded areas mainly differ from natural habitats (usually dominated by <italic><tp:taxon-name>Atriplex portulacoides</tp:taxon-name></italic>, <tp:taxon-name>Chenopodiaceae</tp:taxon-name>, in ungrazed middle marshes) by their enhanced litter layer and by their higher plant cover.</p>
<p>According to <xref ref-type="bibr" rid="B27">McGoech (1998)</xref>, a taxonomic group is an ecological indicator if it responds to environmental changes, stressful or not. In this study, we focussed on ground beetles (<tp:taxon-name>Coleoptera</tp:taxon-name>, <tp:taxon-name>Carabidae</tp:taxon-name>) as they are known to react quickly and strongly to changes in micro-habitat conditions. This group is thus frequently used as an indicator of human disturbances or management practices (e.g., <xref ref-type="bibr" rid="B26">Luff et al. 1992</xref>, <xref ref-type="bibr" rid="B16">Georges 1994</xref>, <xref ref-type="bibr" rid="B41">Sunderland and Lövei 1996</xref>, <xref ref-type="bibr" rid="B37">Rainio and Niemelä 2003</xref>). The assessment of human impact was conducted by comparing two conservation criteria, i.e., abundance of halophilic species and species richness, between natural and invaded stations. Species richness is widely used as a conservation target (e.g., <xref ref-type="bibr" rid="B30">Noss 1990</xref>, <xref ref-type="bibr" rid="B4">Bonn and Gaston 2005</xref>). The use of stenotopic species is also recommended in studying the impact of human activities on arthropod communities (<xref ref-type="bibr" rid="B39">Samways 1993</xref>, <xref ref-type="bibr" rid="B29">New 1995</xref>, <xref ref-type="bibr" rid="B10">Dufrêne and Legendre 1997</xref>). In this study, the target species were halophilic species, defined by their preference or exclusive presence in salt-marsh habitats (<xref ref-type="bibr" rid="B21">Kamer et al. 2008</xref>), which can be assessed using distribution maps (in our study, relevant atlases are <xref ref-type="bibr" rid="B25">Luff 1998</xref> and <xref ref-type="bibr" rid="B46">Turin 2000</xref>). Two approaches were used for assessing changes in natural salt marshes compared to invaded ones: (i) a diachronic (before vs. after the invasion) comparison<!--PageBreak--> of species distribution along a land-sea gradient and (ii) a synchronic comparison of species assemblages between invaded and natural habitats.</p>
</sec>
<sec sec-type="methods"><title>Methods</title>
<sec sec-type="Study site and sampling design"><title>Study site and sampling design</title>
<p>The Mont Saint-Michel bay (NW France) is an extensive littoral zone (500 km²) located between the regions Brittany and Normandy (<named-content content-type="dwc:verbatimCoordinates">48°40’N, 1°40’W</named-content>). Two sites have been studied in salt marshes: “la Ferme Foucault”, on the western part of the Mont St.-Michel (coded F; <named-content content-type="dwc:verbatimCoordinates">48°37’N, 1°32’W</named-content>) and “la Rive” on the eastern part of the Mont St.-Michel (coded R; <named-content content-type="dwc:verbatimCoordinates">48°37’N, 1°29’W</named-content>) (<xref ref-type="fig" rid="F1">Fig. 1</xref>).</p>
<p>For the diachronic approach, ground beetle populations were compared at seven stations (A to G) located along the same land-sea transect at the “Ferme Foucault” site between 1983–1984 and 2002. During the study of 1983–1984, <italic><tp:taxon-name>Elymus athericus</tp:taxon-name></italic> was restricted in this salt marsh to the dyke (station A) and to the upper marsh (station B), but absent from stations C-G. Invasion by <italic><tp:taxon-name>Elymus athericus</tp:taxon-name></italic> modified the plant cover of the sampling stations between 1984 and 2002. The middle marsh and lower marsh stations (station C till F), dominated in 1984 by <italic><tp:taxon-name>Atriplex portulacoides</tp:taxon-name></italic> (<tp:taxon-name>Chenopodiaceae</tp:taxon-name>), were dominated by <italic><tp:taxon-name>Elymus athericus</tp:taxon-name></italic> in 2002.</p>
<p>Secondly, natural (dominated by <italic><tp:taxon-name>Atriplex portulacoides</tp:taxon-name></italic>), and invaded (dominated by <italic><tp:taxon-name>Elymus athericus</tp:taxon-name></italic>) stations were studied at different marsh levels in the synchronic approach. Comparisons of paired stations (natural and invaded – coded N and I, respectively) were spatially replicated three times for avoiding pseudo-replication (<xref ref-type="bibr" rid="B17">Hulbert 1984</xref>). Paired stations were located at the same distance from the dyke because of the existence of a salinity gradient influencing both species richness and abundance (<xref ref-type="bibr" rid="B34">Pétillon et al. 2004</xref>): stations 1 (350m), stations 2 (800–900m; both couples of stations at the “Ferme Foucault” site) and stations 3 (1000 meters from the dyke; “La Rive” site). Because of the clonal progression of the invasive species, all <italic><tp:taxon-name>Elymus</tp:taxon-name></italic> populations (stations I1, I2 and I3) formed a uniform and continuous plant cover. The natural areas sampled were either patch-like formations (in case of stations N1 and N2) or strip-like formations (station N3). Mean salinities did not significantly differ between invaded and natural stations at each salt marsh level (<xref ref-type="bibr" rid="B35">Pétillon et al. 2005</xref>) and elevations were similar between compared stations (J.C. Castel and J. Huet, 1999, unpublished data). More details on the sampling stations can be found in <xref ref-type="bibr" rid="B14">Fouillet (1986)</xref> and <xref ref-type="bibr" rid="B31">Pétillon (2005)</xref>.</p>
</sec>
<sec sec-type="Sampling techniques and species identification"><title>Sampling techniques and species identification</title>
<p>For both the synchronic and diachronic approaches, ground beetles were sampled with pitfall traps, consisting of polypropylene cups (10 cm diameter, 17 cm deep) with <!--PageBreak-->ethylene-glycol as preservative. Traps were covered with a raised wooden roof to keep out rain. They were emptied weekly when tides permitted (i.e., about three weeks per month). Pitfall traps were grouped by four and spaced 10 m apart, this being considered to be the minimum distance for avoiding interference between traps (<xref ref-type="bibr" rid="B44">Topping and Sunderland 1992</xref>). Before the <italic><tp:taxon-name>Elymus</tp:taxon-name></italic> invasion, <xref ref-type="bibr" rid="B14">Fouillet (1986)</xref> sampled the transect with one trap per station from May to September in 1983 and 1984, for a total of 16 five-day samples. In 2002, four traps were installed at each station in both study sites, from April to November 2002. Sampling time was comparable between both periods (90 days in 1983–1984 and 96 days in 2002). Because of the differences in sampling efforts, we only compared the two studies on the basis of species presence / absence (i.e., distribution range along the land-sea transect).</p>
<p>Ground beetles were preserved in 70% ethanol and identified using <xref ref-type="bibr" rid="B20">Jeannel (1942)</xref> and <xref ref-type="bibr" rid="B45">Trautner and Geigenmüller (1987)</xref>. Nomenclature follows <xref ref-type="bibr" rid="B24">Lindroth (1992)</xref> as far as possible, and Fauna Europaea otherwise (http://www.faunaeur.org/).</p>
</sec>
<sec sec-type="Data analyses"><title>Data analyses</title>
<p>Statistics on the abundances of halophilic species were performed only for species represented by at least 10 individuals in couples of stations. Catches in pitfall traps were<!--PageBreak--> related to trapping duration and pitfall trap perimeter, which calculates an “activity trappability density” (number of individuals per day and per meter – <xref ref-type="bibr" rid="B42">Sunderland et al. 1995</xref>). Mean species richness and mean abundances were compared using a two-way mixed model (habitat × station) with habitat type as fixed factor, station (1, 2 and 3) and interaction habitat*station as random factors. In case of non-significant interaction between habitat type and station, the interaction was removed from the model and a new model was performed for detecting significant effects of habitat type and/or station. In case of significant interaction between habitat type and station, parameters were analysed station by station (one-way ANOVA). Statistical analyses were performed using the Statistica-7 software.</p>
<fig id="F1" position="float" orientation="portrait">
<label>Figure 1.</label>
<caption><p>Location of the study sites (Mont St-Michel Bay, France). Codes: <bold>F</bold> ‘Ferme Foucault’ <bold>R</bold> ‘la Rive’.</p></caption>
<graphic xlink:href="ZooKeys-100-407-g001.jpg" position="float" orientation="portrait" xlink:type="simple"/>
</fig>
</sec>
</sec>
<sec sec-type="Results"><title>Results</title>
<sec sec-type="Diachronic approach"><title>Diachronic approach</title>
<p>A total of 24 species (represented by 7,774 individuals) and 35 species (represented by 8,588 individuals) were caught in 1983–1984 and in 2002, respectively. Five species were exclusive to the first sampling period and 16 to the second one. All the species that were only recorded in 1983–1984 were caught in very low numbers (max. 2 individuals), four species on the dyke (<italic><tp:taxon-name>Clivina colaris</tp:taxon-name></italic>, <italic><tp:taxon-name>Dromius linearis</tp:taxon-name></italic>, <italic><tp:taxon-name>Harpalus rufibarbis</tp:taxon-name></italic> and <italic><tp:taxon-name>Harpalus rufipes</tp:taxon-name></italic>) and only one in the salt marsh (<italic><tp:taxon-name>Dyschirius chalceus</tp:taxon-name></italic>). As the sampling effort was quite different between 1983–1984 and 2002 (see Material and Methods), it cannot be concluded that the ‘appearance’ of species between the two studies can be related to the invasion by <italic><tp:taxon-name>Elymus athericus</tp:taxon-name></italic>. The comparison in distribution was thus restricted to the 19 shared species (<xref ref-type="table" rid="T1">Table 1</xref>).</p>
<p>In terms of distribution ranges, two groups of carabids were distinguished: species with constant distribution range in the salt marsh or on the dyke and species with an increased distribution range between 1983–1984 and 2002. Eight species were caught on the dyke in 1983–1984 and in 2002, and seemed not to have progressed with <italic><tp:taxon-name>Elymus athericus</tp:taxon-name></italic> in the salt marsh (<italic><tp:taxon-name>Amara equestris</tp:taxon-name></italic>, <italic><tp:taxon-name>Anisodactylus binotatus</tp:taxon-name></italic>, <italic><tp:taxon-name>Bembidion tetracolum</tp:taxon-name></italic>, <italic><tp:taxon-name>Harpalus anxius</tp:taxon-name></italic>, <italic><tp:taxon-name>Leistus fulvibarbis</tp:taxon-name></italic>, <italic><tp:taxon-name>Nebria brevicollis</tp:taxon-name></italic>, <italic><tp:taxon-name>Pterostichus melanarius</tp:taxon-name></italic> and <italic><tp:taxon-name>Pterostichus niger</tp:taxon-name></italic>: <xref ref-type="table" rid="T1">Table 1</xref>). Eight other species had a similar habitat range in the salt marsh, extending from the upper to lower marsh or from the dyke to the lower marsh (halophilic species: bold in <xref ref-type="table" rid="T1">Table 1</xref>), plus two high-marsh living species (<italic><tp:taxon-name>Badister bipustulatus</tp:taxon-name></italic> and <italic><tp:taxon-name>Pterostichus vernalis</tp:taxon-name></italic>), one low-marsh living species (<italic><tp:taxon-name>Dyschirius salinus</tp:taxon-name></italic>) and one species with a discontinuous range along the land-sea transect (<italic><tp:taxon-name>Loricera pilicornis</tp:taxon-name></italic>).Only three speciesshowed an extension of their distribution in the salt marsh, both to the upper and lower marsh (<italic><tp:taxon-name>Bembidion iricolor</tp:taxon-name></italic>, <italic><tp:taxon-name>Bembidion lampros</tp:taxon-name></italic> and <italic><tp:taxon-name>Pterostichus cupreus</tp:taxon-name></italic>).</p>
<p>The <italic><tp:taxon-name>Elymus athericus</tp:taxon-name></italic> invasion led to a decrease in the percentage of halophilic species in invaded salt marshes (<xref ref-type="fig" rid="F2">Fig. 2</xref>).</p>
<fig id="F2" position="float" orientation="portrait">
<label>Figure 2.</label>
<caption><p>Changes in the percentage of halophilic species in the salt marsh after the invasion by <italic><tp:taxon-name>Elymus athericus</tp:taxon-name></italic>.</p></caption>
<graphic xlink:href="ZooKeys-100-407-g002.jpg" position="float" orientation="portrait" xlink:type="simple"/>
</fig>
<table-wrap position="float" orientation="portrait" id="T1">
<label>Table 1.</label>
<caption><p>Comparison of total catches (number of individuals) between 1983–1984 and 2002 along a land-sea transect (Foucault site; bold: halophilic species). The letters <bold>A–G</bold> indicate different sampling stations. In 1983–1984, only stations <bold>A–B</bold> had a dominant <italic><tp:taxon-name>Elymus athericus</tp:taxon-name></italic> cover; in 2002 at all stations <italic><tp:taxon-name>Elymus athericus</tp:taxon-name></italic> was present (dominant cover for stations <bold>A</bold> to <bold>F</bold>).</p></caption>
<table>
<tbody>
<tr>
<th rowspan="1" colspan="1"/>
<th rowspan="1" colspan="1"><italic>Period</italic></th>
<th rowspan="1" colspan="1"><italic>A</italic></th>
<th rowspan="1" colspan="1"><italic>B</italic></th>
<th rowspan="1" colspan="1"><italic>C</italic></th>
<th rowspan="1" colspan="1"><italic>D</italic></th>
<th rowspan="1" colspan="1"><italic>E</italic></th>
<th rowspan="1" colspan="1"><italic>F</italic></th>
<th rowspan="1" colspan="1"><italic>G</italic></th>
<th rowspan="1" colspan="1"><italic>Total</italic></th>
</tr>
<tr>
<td rowspan="1" colspan="10"><italic>SHARED SPECIES</italic></td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Amara equestris</tp:taxon-name></italic>(Duftschmid 1812)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Anisodactylus binotatus</tp:taxon-name></italic>(Fabricius 1787)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">3</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">3</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">6</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">6</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Badister bipustulatus</tp:taxon-name></italic>(Fabricius 1792)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">4</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Bembidion iricolor</tp:taxon-name></italic>Bedel 1879</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">6</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">5</td>
<td rowspan="1" colspan="1">98</td>
<td rowspan="1" colspan="1">89</td>
<td rowspan="1" colspan="1">21</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1">4</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">218</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Bembidion lampros</tp:taxon-name></italic>(Herbst 1784)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">3</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1">12</td>
<td rowspan="1" colspan="1">18</td>
<td rowspan="1" colspan="1">3</td>
<td rowspan="1" colspan="1">5</td>
<td rowspan="1" colspan="1">4</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1">44</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Bembidion minimum</tp:taxon-name></italic> (Fabricius 1792)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">4</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1">40</td>
<td rowspan="1" colspan="1">52</td>
<td rowspan="1" colspan="1">10</td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">109</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1">31</td>
<td rowspan="1" colspan="1">13</td>
<td rowspan="1" colspan="1">5</td>
<td rowspan="1" colspan="1">80</td>
<td rowspan="1" colspan="1">96</td>
<td rowspan="1" colspan="1">3</td>
<td rowspan="1" colspan="1">229</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Bembidion normanum</tp:taxon-name></italic> Dejean 1831</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">24</td>
<td rowspan="1" colspan="1">39</td>
<td rowspan="1" colspan="1">244</td>
<td rowspan="1" colspan="1">149</td>
<td rowspan="1" colspan="1">53</td>
<td rowspan="1" colspan="1">510</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1">6</td>
<td rowspan="1" colspan="1">8</td>
<td rowspan="1" colspan="1">13</td>
<td rowspan="1" colspan="1">212</td>
<td rowspan="1" colspan="1">131</td>
<td rowspan="1" colspan="1">24</td>
<td rowspan="1" colspan="1">396</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Bembidion tetracolum</tp:taxon-name></italic> (Say 1823)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Dicheirotrichus gustavii</tp:taxon-name></italic> Crotch 1871</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">11</td>
<td rowspan="1" colspan="1">83</td>
<td rowspan="1" colspan="1">2121</td>
<td rowspan="1" colspan="1">2622</td>
<td rowspan="1" colspan="1">393</td>
<td rowspan="1" colspan="1">5232</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1">8</td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1">3</td>
<td rowspan="1" colspan="1">136</td>
<td rowspan="1" colspan="1">237</td>
<td rowspan="1" colspan="1">156</td>
<td rowspan="1" colspan="1">544</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Dyschirius salinus</tp:taxon-name></italic> Schaum 1843</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1">2</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">5</td>
<td rowspan="1" colspan="1">5</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Harpalus anxius</tp:taxon-name></italic>(Duftschmid 1812)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">2</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Leistus fulvibarbis</tp:taxon-name></italic> Dejean 1826</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">3</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">3</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Loricera pilicornis</tp:taxon-name></italic>(Fabricius 1775)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1">2</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Nebria brevicollis</tp:taxon-name></italic> (Fabricius 1792)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">3</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">3</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Pogonus chalceus</tp:taxon-name></italic> (Marsham 1802)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">8</td>
<td rowspan="1" colspan="1">4</td>
<td rowspan="1" colspan="1">65</td>
<td rowspan="1" colspan="1">42</td>
<td rowspan="1" colspan="1">678</td>
<td rowspan="1" colspan="1">617</td>
<td rowspan="1" colspan="1">436</td>
<td rowspan="1" colspan="1">1850</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">13</td>
<td rowspan="1" colspan="1">100</td>
<td rowspan="1" colspan="1">193</td>
<td rowspan="1" colspan="1">126</td>
<td rowspan="1" colspan="1">1628</td>
<td rowspan="1" colspan="1">1290</td>
<td rowspan="1" colspan="1">2243</td>
<td rowspan="1" colspan="1">5593</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Pterostichus cupreus</tp:taxon-name></italic>(Linnaeus 1758)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">3</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">5</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">8</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">7</td>
<td rowspan="1" colspan="1">41</td>
<td rowspan="1" colspan="1">9</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">59</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Pterostichus niger</tp:taxon-name></italic>(Schaller 1783)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">24</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">24</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Pterostichus vernalis</tp:taxon-name></italic>(Panzer 1795)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">4</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">4</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">3</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Pterostichus melanarius</tp:taxon-name></italic>(Illiger 1798)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">4</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">4</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">12</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">12</td>
</tr>
<tr>
<td rowspan="1" colspan="10"><italic>SPECIES NOT RECOLLECTED IN 2002</italic></td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Clivina collaris</tp:taxon-name></italic>(Herbst 1786)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">2</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Dromius linearis</tp:taxon-name></italic>(Olivier 1795)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Dyschirius chalceus</tp:taxon-name></italic> Erichson 1837</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Harpalus rufibarbis</tp:taxon-name></italic>(Fabricius 1792)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">2</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Harpalus rufipes</tp:taxon-name></italic>(Degeer 1774)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td rowspan="1" colspan="10"><italic>NEW SPECIES FOUND IN 2002</italic></td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Anchomenus dorsalis</tp:taxon-name></italic>(Pontoppidan 1763)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Agonum muelleri</tp:taxon-name></italic>(Herbst 1784)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">5</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">6</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Amara lunicollis</tp:taxon-name></italic>Schiödte 1837</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">3</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">3</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Amara plebeja</tp:taxon-name></italic>(Gyllenhal 1810)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1">3</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Amara tibialis</tp:taxon-name></italic>(Paykull 1798)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">2</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Anisodactylus poeciloides</tp:taxon-name></italic> (Stephens 1828)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">2</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Bembidion obtusum</tp:taxon-name></italic>Serville 1821</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">10</td>
<td rowspan="1" colspan="1">16</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">27</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Calathus mollis</tp:taxon-name></italic>(Marsham 1802)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Clivina fossor</tp:taxon-name></italic>(Linnaeus 1758)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Dicheirotrichus obsoletus</tp:taxon-name></italic> (Dejean 1829)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1">12</td>
<td rowspan="1" colspan="1">5</td>
<td rowspan="1" colspan="1">478</td>
<td rowspan="1" colspan="1">572</td>
<td rowspan="1" colspan="1">301</td>
<td rowspan="1" colspan="1">1370</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Harpalus distinguendus</tp:taxon-name></italic>(Duftschmid 1812)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">3</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Harpalus melancholichus</tp:taxon-name></italic>Dejean 1829</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td rowspan="1" colspan="1">2002</td>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">1</td>
</tr>
<tr>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Microlestes minutulus</tp:taxon-name></italic>(Goeze 1777)</td>
<td rowspan="1" colspan="1">1983–84</td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1"> </td>
<td rowspan="1" colspan="1">0</td>