639:
14.6 ± 4.8 °C (58.3 ± 8.6 °F), slightly higher than seen at
Republic, and CLAMP analysis gave an overall mean annual temperature of 13.3 ± 2.1 °C (55.9 ± 3.8 °F). A bioclimatic-based estimate based on modern relatives of the taxa found at each site suggested mean annual temperatures around 13.5 ± 2.2 °C (56.3 ± 4.0 °F) for Republic and 14.7 ± 2.1 °C (58.5 ± 3.8 °F) for Quilchena. These are lower than the mean annual temperature estimates given for the coastal Puget Group, which is estimated to have been between 15–18.6 °C (59.0–65.5 °F). The bioclimatic analysis for Republic and Falkland suggest mean annual precipitation amounts of 115 ± 39 cm (45 ± 15 in) and 121 ± 39 cm (48 ± 15 in) respectively.
275:
125:
104:
622:
range between 0.7–1.2 km (0.43–0.75 mi) higher than the coastal forests. This is consistent with the paleoelevation estimates for the lake systems, which range between 1.1–2.9 km (1,100–2,900 m), which is similar to the modern elevation 0.8 km (0.50 mi), but higher.
638:
of both paleofloras. The CLAMP results after multiple linear regressions for
Republic gave a mean annual temperature of approximately 8.0 °C (46.4 °F), with the LMA giving 9.2 ± 2.0 °C (48.6 ± 3.6 °F). LMA results from Quilchena returned the higher
573:
separating them. The inner most set of lateral primary veins run parallel to the median secondary veins, broadly curving upwards and with three secondary veins branching off the exterior side. The branched secondaries run parallel to the next lateral primary vein. The tertiary veins run
568:
is palmate in venation with an overall orbicular shape, cordate blade base and acute blade tip. The central primary vein is flanked by three pairs of lateral primary veins and the margin of the leaf has evenly spaced, distinctly shaped teeth with rounded
582:
Formations of the
Okanagan Highlands formations represent upland lake systems that were surrounded by a warm temperate ecosystem with nearby volcanism dating from during and just after the early Eocene climatic optimum. The highlands likely had a
696:
Greenwood, D.R.; Archibald, S.B.; Mathewes, R.W; Moss, P.T. (2005). "Fossil biotas from the
Okanagan Highlands, southern British Columbia and northeastern Washington State: climates and ecosystems across an Eocene landscape".
815:
Rubino, E.; Leier, A.; Cassel, E.; Archibald, S.; Foster-Baril, Z.; Barbeau, D. Jr (2021). "Detrital zircon UPb ages and Hf-isotopes from Eocene intermontane basin deposits of the southern
Canadian Cordillera".
594:
climate, in which winter temperatures rarely dropped low enough for snow, and which were seasonably equitable. The paleoforest surrounding the lakes have been described as precursors to the modern
735:
Moss, PT; Greenwood, DR; Archibald, SB (2005). "Regional and local vegetation community dynamics of the Eocene
Okanagan Highlands (British Columbia – Washington State) from palynology".
574:
perpendicular to the secondary veins with an even spacing, while the quaternary veins are orthogonal to the tertiaries forming reticulated pattern of pentagonal and quadrangular spaces.
770:
Makarkin, V.; Archibald, S.B. (2014). "An unusual new fossil genus probably belonging to the
Psychopsidae (Neuroptera) from the Eocene Okanagan Highlands, western North America".
1004:
491:
459:, and is reconstructed as the warmest and wettest of the Early Eocene upland sites from the Okanagan Highlands of British Columbia and northern Washington State.
989:
994:
1014:
595:
1029:
1009:
999:
541:
1019:
549:
385:
515:
548:
was the oldest macrofossil occurrence for the genus to be described,; older microfossil records of pollen date near to the
420:
396:
microfossils and macrofossils have not been identified from the
Horsefly fossil beds near the unincorporated community of
381:
498:. Working from this specimen, collected in the Republic, Washington area in the early 1980s, the fossil was studied by
365:
341:
274:
397:
389:
377:
369:
357:
349:
337:
862:
533:
503:
124:
412:
914:
779:
961:
242:
877:
863:"Paleoenvironment of the Quilchena flora, British Columbia, during the Early Eocene Climatic Optimum"
744:
706:
345:
784:
952:
611:
408:
839:
1. Early Eocene
Lagerstätten of the Okanagan Highlands (British Columbia and Washington State)".
487:
424:
373:
318:
119:
966:
797:
361:
893:
885:
835:
Archibald, S.; Greenwood, D.; Smith, R.; Mathewes, R.; Basinger, J. (2011). "Great
Canadian
789:
752:
714:
635:
631:
322:
615:
292:
162:
423:(EECO). Modern work on the fossil-bearing strata of the Klondike Mountain Formation via
881:
748:
710:
1024:
570:
522:
511:
507:
149:
983:
599:
584:
499:
471:
261:
61:
587:
483:
404:
946:
607:
591:
537:
479:
265:
103:
36:
937:
793:
495:
81:
46:
663:
664:"Middle Eocene dicotyledonous plants from Republic, northeastern Washington"
619:
518:
296:
208:
86:
889:
801:
931:
529:
475:
446:
428:
416:
306:
198:
175:
76:
71:
56:
51:
41:
627:
288:
91:
66:
898:
454:
440:
433:
603:
452:
isotopic data published in 2021. The Quilchena locality is dated to
449:
314:
310:
188:
908:
756:
718:
301:
273:
218:
136:
912:
598:
of Eastern North America and Eastern Asia. Based on the fossil
305:, is related to modern lindens (called "limes" in Britain and "
861:
Mathewes, R. W.; Greenwood, D. R.; Archibald, S. B. (2016).
532:
recognizing the help provided to Wolfe and Wehr by a young
336:
leaf fossils have been identified from two locations in the
544:. Wolfe and Wehr noted that, at the time of publication,
403:
The age for the Okanagan Highland locations is uniformly
521:
on the North Eastern Washington dicot fossils. The
921:
632:climate leaf analysis multivariate program (CLAMP)
510:of the Burke Museum. They published their 1987
321:, United States and a similar aged formation in
856:
854:
691:
689:
687:
685:
683:
681:
556:relative are known from the Eocene of England.
492:University of California Museum of Paleontology
438:at the youngest, and an oldest age estimate of
614:of Western Washington, which are described as
352:. Fossil pollen identified as from the genus
8:
1005:Flora of the West Coast of the United States
457: ± 0.4 million years ago
443: ± 0.1 million years ago
356:has been identified from a greater range of
436: ± .5 million years ago
909:
730:
728:
602:the lakes were higher and cooler then the
102:
20:
897:
783:
657:
655:
653:
651:
634:analysis of the Republic paleoflora, and
309:" in the US). The species is known from
668:United States Geological Survey Bulletin
647:
427:has given an estimated age in the Late
596:temperate broadleaf and mixed forests
7:
392:. Of the Okanagan Highlands sites,
431:stage of the early Eocene, between
407:, with the sites that have current
368:, at the Falkland fossil site near
348:and at the Quilchena locality near
870:Canadian Journal of Earth Sciences
737:Canadian Journal of Earth Sciences
699:Canadian Journal of Earth Sciences
542:National Museum of Natural History
16:Extinct species of flowering plant
14:
606:coastal forests preserved in the
990:Ypresian plants of North America
662:Wolfe, J.A.; Wehr, W.C. (1987).
386:Driftwood Canyon Provincial Park
123:
516:United States Geological Survey
299:that, as a member of the genus
995:Extinct flora of North America
358:Okanagan Highland fossil sites
1:
1015:Fossil taxa described in 1987
618:ecosystems. Estimates of the
421:Early Eocene Climatic Optimum
419:age and corresponding to the
470:was described from a single
1030:Klondike Mountain Formation
1010:Flora of Washington (state)
552:, and fruits of an extinct
550:Paleocene – Eocene boundary
415:radiometric dates being of
366:Princeton, British Columbia
360:, having been found in the
342:Klondike Mountain Formation
1046:
636:leaf margin analysis (LMA)
463:History and classification
313:leaves found in the early
1000:Flora of Northern America
794:10.11646/zootaxa.3838.3.8
350:Merritt, British Columbia
338:Eocene Okanagan Highlands
248:
241:
120:Scientific classification
118:
110:
101:
23:
1020:Plants described in 1987
504:University of California
630:have been derived from
628:mean annual temperature
616:lowland tropical forest
478:being UW 39712, in the
278:Holotype with herbivory
890:10.1139/cjes-2015-0163
536:, now director of the
425:radiometrically dating
279:
962:Paleobiology Database
514:for the species in a
317:deposits of northern
277:
346:Republic, Washington
882:2016CaJES..53..574M
818:Sedimentary Geology
749:2005CaJES..42..187M
711:2005CaJES..42..167G
612:Chuckanut Formation
588:upper microthermal
374:McAbee Fossil Beds
280:
977:
976:
915:Taxon identifiers
841:Geoscience Canada
626:Estimates of the
592:lower mesothermal
564:The type leaf of
490:UCMP 9291 in the
445:, given based on
362:Allenby Formation
272:
271:
1037:
970:
969:
957:
956:
955:
942:
941:
940:
910:
904:
903:
901:
867:
858:
849:
848:
832:
826:
825:
812:
806:
805:
787:
767:
761:
760:
732:
723:
722:
693:
676:
675:
659:
578:Paleoenvironment
523:specific epithet
512:type description
458:
444:
437:
323:British Columbia
319:Washington state
257:
253:
234:T. johnsoni
128:
127:
106:
96:
33:
29:Temporal range:
21:
1045:
1044:
1040:
1039:
1038:
1036:
1035:
1034:
980:
979:
978:
973:
965:
960:
951:
950:
945:
936:
935:
930:
923:Tilia johnsonii
917:
907:
865:
860:
859:
852:
834:
833:
829:
814:
813:
809:
785:10.1.1.692.1185
769:
768:
764:
757:10.1139/e04-095
734:
733:
726:
719:10.1139/e04-100
695:
694:
679:
661:
660:
649:
645:
580:
562:
482:collections of
465:
453:
439:
432:
382:Hat Creek Amber
331:
293:flowering plant
268:
259:
251:
250:
237:
122:
111:Fossil leaf of
97:
95:
94:
89:
84:
79:
74:
69:
64:
59:
54:
49:
44:
39:
31:
30:
27:
17:
12:
11:
5:
1043:
1041:
1033:
1032:
1027:
1022:
1017:
1012:
1007:
1002:
997:
992:
982:
981:
975:
974:
972:
971:
958:
953:Tilia johnsoni
943:
927:
925:
919:
918:
913:
906:
905:
876:(6): 574–590.
850:
827:
807:
778:(3): 385–391.
762:
743:(2): 187–204.
724:
705:(2): 167–185.
677:
646:
644:
641:
620:paleoelevation
579:
576:
566:Tilia johnsoni
561:
558:
508:Wesley C. Wehr
480:paleobotanical
474:, a leaf, the
468:Tilia johnsoni
464:
461:
334:Tilia johnsoni
330:
327:
295:in the family
284:Tilia johnsoni
270:
269:
260:
255:Tilia johnsoni
246:
245:
239:
238:
230:
228:
224:
223:
216:
212:
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206:
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196:
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116:
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99:
98:
90:
85:
80:
75:
70:
65:
60:
55:
50:
45:
40:
35:
34:
28:
25:Tilia johnsoni
15:
13:
10:
9:
6:
4:
3:
2:
1042:
1031:
1028:
1026:
1023:
1021:
1018:
1016:
1013:
1011:
1008:
1006:
1003:
1001:
998:
996:
993:
991:
988:
987:
985:
968:
963:
959:
954:
948:
944:
939:
933:
929:
928:
926:
924:
920:
916:
911:
900:
895:
891:
887:
883:
879:
875:
871:
864:
857:
855:
851:
847:(4): 155–164.
846:
842:
838:
831:
828:
823:
819:
811:
808:
803:
799:
795:
791:
786:
781:
777:
773:
766:
763:
758:
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750:
746:
742:
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716:
712:
708:
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678:
673:
669:
665:
658:
656:
654:
652:
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642:
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633:
629:
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621:
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613:
609:
605:
601:
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572:
567:
559:
557:
555:
551:
547:
543:
539:
535:
531:
527:
524:
520:
517:
513:
509:
505:
501:
500:Jack A. Wolfe
497:
493:
489:
485:
481:
477:
473:
472:type specimen
469:
462:
460:
456:
451:
448:
442:
435:
430:
426:
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418:
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406:
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379:
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367:
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328:
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324:
320:
316:
312:
308:
304:
303:
298:
294:
290:
286:
285:
276:
267:
263:
258:
256:
247:
244:
243:Binomial name
240:
236:
235:
229:
226:
225:
222:
221:
217:
214:
213:
210:
207:
204:
203:
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184:
181:
180:
177:
174:
171:
168:
167:
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161:
158:
155:
154:
151:
150:Tracheophytes
148:
145:
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141:
138:
135:
132:
131:
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117:
114:
109:
105:
100:
93:
88:
83:
78:
73:
68:
63:
58:
53:
48:
43:
38:
26:
22:
19:
922:
873:
869:
844:
840:
837:Lagerstätten
836:
830:
821:
817:
810:
775:
771:
765:
740:
736:
702:
698:
671:
667:
625:
581:
565:
563:
553:
545:
534:Kirk Johnson
525:
484:Burke Museum
467:
466:
409:uranium-lead
405:Early Eocene
402:
393:
353:
333:
332:
329:Distribution
300:
283:
282:
281:
254:
249:
233:
232:
219:
182:
169:
156:
143:
112:
24:
18:
947:Wikispecies
608:Puget Group
560:Description
546:T. johnsoni
538:Smithsonian
488:counterpart
413:argon–argon
291:species of
163:Angiosperms
113:T. johnsoni
984:Categories
899:1807/71979
643:References
496:California
486:, and its
325:, Canada.
824:(online).
780:CiteSeerX
519:monograph
307:basswoods
297:Malvaceae
227:Species:
209:Malvaceae
133:Kingdom:
938:Q7802148
932:Wikidata
802:25081783
530:patronym
526:johnsoni
476:holotype
447:detrital
429:Ypresian
417:Ypresian
398:Horsefly
390:Smithers
378:Kamloops
370:Falkland
205:Family:
199:Malvales
176:Eudicots
32:Ypresian
878:Bibcode
772:Zootaxa
745:Bibcode
707:Bibcode
674:: 1–25.
571:sinuses
502:of the
289:extinct
252:†
231:†
215:Genus:
195:Order:
137:Plantae
967:463626
822:105969
800:
782:
604:coeval
600:biotas
450:zircon
380:, the
372:, the
340:, the
315:Eocene
311:fossil
287:is an
264:&
189:Rosids
1025:Tilia
866:(PDF)
585:mesic
554:Tilia
528:is a
394:Tilia
388:near
376:near
364:near
354:Tilia
344:near
302:Tilia
262:Wolfe
220:Tilia
183:Clade
170:Clade
157:Clade
144:Clade
798:PMID
776:3838
672:1597
610:and
506:and
455:51.5
441:51.2
434:49.4
384:and
266:Wehr
37:PreꞒ
894:hdl
886:doi
790:doi
753:doi
715:doi
590:to
540:'s
494:in
411:or
986::
964::
949::
934::
892:.
884:.
874:53
872:.
868:.
853:^
845:38
843:.
820:.
796:.
788:.
774:.
751:.
741:42
739:.
727:^
713:.
703:42
701:.
680:^
670:.
666:.
650:^
400:.
185::
172::
159::
146::
87:Pg
902:.
896::
888::
880::
804:.
792::
759:.
755::
747::
721:.
717::
709::
92:N
82:K
77:J
72:T
67:P
62:C
57:D
52:S
47:O
42:Ꞓ
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