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Herpetological assemblages of the Michigan Regional Landscape Ecosystems.

ABSTRACT

The herpetofauna of each of the four Michigan Regional Landscape Ecosystems (Region I, southern Lower Peninsula; II, northern LP; III, eastern Upper Peninsula; IV, western Upper Peninsula) was determined based on museum specimens and records from ongoing studies. These regional herpetofaunas were compared on the basis of (1) taxonomic diversity at the species level; (2) presence of Primary, Secondary, and Tertiary re-entrant post-glacial taxa; (3) presence of gaps in the distribution of widespread regional taxa; and (4) presence of relict species. Region I is the most taxonomically diverse having 94.4% of the species that occur in Michigan. Region II is somewhat less diverse having 90.2% of I. Regions III and IV are much less specifically diverse than I and II; III having 64.7% and IV having 66.7% of I. The main difference comes from the fact that reptiles are much less specifically diverse in III and IV than in I and II--where II has 93.1%, III only 51.7%, and IV only 55.2% of I. Amphibian diversity is relatively constant in all three northern regions where II has 86.4%, III 81.8% and IV 81.8% of I. Region I has six Tertiary re-entrant taxa but II, III, and IV have none of these. Region I has significantly more gaps in the distribution of generally widespread taxa (20 species) than the other regions. Region I has five relict species, II has six, III has five, and IV has three. These relict populations may be mainly due to a significant warm, dry spell (hypsithermal) in the mid-Holocene and a significant cold spell (Little Ice Age) at the end of the Holocene.

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INTRODUCTION

Major herpetological treatises in the Great Lakes states (e.g., Illinois, Smith 1961; Indiana, Minton 2001; Ohio, Conant 1951; Wisconsin, Vogt 1981) have, in part, detailed regional variations in the herpetofaunas of their respective states. But such a treatise in herpetology for Michigan has not been done since Ruthven et al. (1928), who reported only a part of the species that are now known in the state (see Conant and Collins 1998; Harding 1997; Harding and Holman 1990, 1992; Holman 1994, 2001a,b; and Holman et al. 1993). Moreover, Ruthven et al. did not discuss regional differences in the Michigan herpetofauna. The present paper attempts to rectify this situation by a report and discussion on the herpetofauna of each of the four Michigan Regional Landscape Ecosystems (MRLE).

The four regional landscape ecosystems in Michigan have been most recently discussed by Barnes and Wagner (2004) and were previously discussed by Albert (1995), Denton and Barnes (1988), Dickmann and Leefers (2003), Edgar (1991), and Holman and Holman (2003). These four regional systems are based on the combined effect of (1) glacially derived landforms; (2) underlying bedrock; (3) soils; and (4) climate (Dickmann and Leefers 2003).

I feel that the regional ecosystem approach provides a more logical basis for the analysis of the Michigan herpetofauna than previous widely used "natural" subdivisions. The North American Biotic Provinces system of Dice (1943) divides Michigan into a southern (Carolinian) province and a northern (Canadian) one based on dominant vegetational and mammalian assemblages (later a transition zone between the two was recognized, e.g., Holman and Holman 2003). The so-called Mason-Quimby line (see Holman 1991, 2001c, and Holman and Holman 2003) also divides Michigan into a northern and a southern region (see Holman 2001c, Figure 10, 19). This line is based on the lack of mastodont, mammoth, and other Pleistocene vertebrate remains as well as Paleo-Indian artifacts north of the line.

A recent survey by Holman (2001a) showed, not unexpectedly, that the distribution of many amphibians and reptiles in Michigan is much more fragmentary and spotty than modern range maps (especially those in field guides) indicate. This led to a publication on the herpetofauna of the fossil dunes and soils near Saginaw Bay (Holman 2001b), where it was suggested that the area was a unique herpetological habitat. The present paper is, in part, an outgrowth of that publication.

The condition of the modern herpetofauna in Michigan is undoubtedly due, in part, to human modification of the environment. But beyond this, herpetological habitats were quite recently obliterated by Pleistocene ice masses and then re-occupied again, probably in a relatively random fashion, rather than by an orderly march of plant and animal species northward. Moreover, some Michigan herpetological species are undoubtedly in the process of re-occupying new habitats (Holman 1992) while others have been left as relicts since the Middle Holocene warm, dry spell (hypsithermal event) was replaced by a cooler climate, especially the "Little Ice Age" that occurred only a few hundred years ago (Bernabo 1981; Kapp 1999; Holman et al. 2003).

THE FOUR MICHIGAN REGIONAL LANDSCAPE ECOSYSTEMS

Regions I and II are in the Lower Peninsula and regions III and IV are in the Upper Peninsula of Michigan (Figure 1). These descriptions are modified from Barnes and Wagner (2004), Dickmann and Leefers (2003), and Edgar (1991).

Region I. This region is warmer throughout the year than the other three and has a plant growing season that is less changeable than the others. Extensive lake plains of sand and clay and ground and end moraines as well as other glacial end features are characteristic of this area. The ridges in I are lower than those to the north and range from about 80 to 160 kilometers high. The dominant soils are loams and clays and sands are less common. The vegetation in Region I includes many southern plants at the northern limit of their range, resulting in plant communities that are more diverse than in the northern regions. The region is dominated by deciduous hardwood (broadleaf) tree species. Natural prairies and savannas formerly were found in much of this region, but have been mainly lost to urban and agricultural expansion. Many animal species that occur in Region I are absent in Regions II, III, and IV.

[FIGURE 1 OMITTED]

Region II. Region II has a much different climate than Region I resulting from (1) being in a more northern latitude; (2) being surrounded on three sides by the Great Lakes; and (3) having uplands that are not only more extensive, but higher than those in I. These upland areas contribute to a cooler, more variable climate with a more likely chance of frost during the plant growing season. Precipitation is influenced in II not only by the more extensive, high uplands, but by the surrounding lakes as well. Sandy soils are more abundant in Region II and extensive glacial deposits are abundant. A dominant feature of II is the presence of forests of pure conifer or mixed conifer/broadleaf forests. Conifers tend to be dominant in the extensive areas of upland, sandy, outwash plains in II, whereas hardwoods tend to dominate the upper slopes of moraines and other high topography. Lowland swamps and bogs are more common and often more acidic than those of Region I. Both plant and animal associations are less diverse in II than in I. [Note: Here Beaver and Bois Blanc Islands are considered part of Region II].

Region III. This region, which comprises the eastern part of the Upper Peninsula, features relatively low elevation and relatively young Paleozoic bedrock formed of patches of Devonian rocks just north of the Straits and more extensive Silurian and Ordovician rocks beyond. This bedrock occurs relatively near the surface in some areas. The plant growing season of 120 to 140 days is similar to that in Region II. The three Great Lakes that surround Region III tend to modify extremes in temperatures. The soils of this region are poor and consist mainly of poorly drained sands and clays. Northern coniferous trees occur in the extensive low places, especially in the characteristic swamp coniferous forests of the extensive wetlands of the area. Upland areas are dominated by mixed conifer/broadleaf forest. [Note: Here Drummond Island is considered a part of Region III.]

Region IV. Region IV comprises the western part of the Upper Peninsula. Here Cambrian rocks and a very complex system of Precambrian rocks form the important bedrock structures of the area. Frequent outcrops of these rocks occur and many uplands and even low mountains are present. This upland topography controls the vegetation of the region and the growing season of plants may be as short as 60 days. Region IV has the most continental climate of the four Michigan Regional Landscape Ecosystems, mainly because there is less lake moderation. The climate is very cold in the winter and warm, in fact sometimes hot, during the growing season. Northern conifer/broadleaf forest associations are abundant, and white pine, red pine, and oak are dominant in the upland forests. [Note: Here Isle Royale is considered part of Region IV.]

MATERIALS AND METHODS

Since 1991 I have attempted to document the herpetofauna of Michigan by checking as many museum records as possible as well as records documented by modern herpetologists active in Michigan. Museums include the American Museum of Natural History, the Field Museum of Natural History, the Illinois Natural History Survey, the Milwaukee Public Museum, the Michigan State University Museum, and the University of Michigan Museum of Zoology. The works of Pentecost and Vogt (1976) and Casper (2002) were consulted.

James H. Harding of the Michigan State University Museum and Lori G. Sargent of the Michigan Department of Natural Resources (MDNR) provided new herpetological records from their ongoing research projects in Michigan. The Michigan Natural Faunal Inventory of the MDNR also provided new records of amphibians and reptiles from the state. The University of Michigan Museum of Zoology Program online on the Distribution of Amphibians and Reptiles in Michigan produced by Greg Schneider and Arnold Kluge was very helpful.

The maps used here (Appendix) indicate county occurrences for each species rather than dots reflecting localities where each species was collected (e.g., as in Minton 2001). Michigan's herpetofauna has already been targeted by illegal collectors in the pet trade (James H. Harding, personal communications) and dots on maps can be used by these individuals in an attempt to find local herpetological "hot spots." This can be especially devastating to rare and endangered species that usually occur in isolated populations; but this practice can also damage so-called "common species," many of which also occur in isolated habitats. Scientific and common names are based on Crother (2000) and Crother et al. (2003).

RESULTS

The MRLE Distribution of the Michigan Herpetofauna

The Michigan herpetofauna is listed below with the MRLE (Roman numerals in parentheses) listed after each taxon. This list contains both species and subspecific designations as well as information about intergrading populations.

Amphibians (I, II, III, IV):

Caudata--Salamanders (I, II, III, IV)

Family Ambystomatidae (I, II, III, IV)

Ambystoma laterale Hallowell, 1856, Blue-spotted Salamander (I, II, III, IV)

Ambystoma maculatum (Shaw, 1802), Spotted Salamander (I, II, III, IV)

Ambystoma opacum (Gravenhorst, 1807), Marbled Salamander (I)

Ambystoma texanum (Matthes, 1855), Small-mouthed Salamander (I)

Ambystoma tigrinum tigrinum (Green, 1825), Eastern Tiger Salamander (I, II, III)

Family Plethodontidae (I, II, III, IV)

Hemidactylium scutatum (Temminck and Schlegel in Von Siebold, 1838), Four-toed Salamander (I, II, III, IV)

Plethodon cinereus (Green, 1818), Eastern Red-backed Salamander (I, II, III, IV)

Family Proteidae (I, II, III, IV)

Necturus maculosus maculosus (Rafinesque, 1818), Common Mudpuppy (I, II, III, IV)

Family Salamandridae (I, II, III, IV)

Notophthalmus viridescens louisianensis Wolterstorff, 1914, Central Newt (I, II, III, IV)

Notopthalmus viridescens viridescens (Rafinesque, 1820), Red-spotted Newt (I)

Family Sirenidae (I)

Siren intermedia nettingi Goin, 1942, Western Lesser Siren (I)

Anura--Frogs and Toads (I, II, III, IV)

Family Bufonidae (I, II, III, IV)

Bufo americanus americanus Holbrook, 1836, Eastern American Toad (I, II, III, IV)

Bufo fowleri Hinckley, 1882, Fowler's Toad (I, II)

Family Hylidae (I, II, III, IV)

Acris crepitans blanchardi Harper, 1947, Blanchard's Cricket Frog (I, II)

Hyla chrysoscelis Cope, 1880, Cope's Gray Treefrog (I, II, III, IV)

Hyla versicolor LeConte, 1825, Gray Treefrog (I, II, III, IV)

Pseudacris crucifer crucifer (Wied-Neuwied, 1838), Northern Spring Peeper (I, II, III, IV)

Pseudacris maculata (Agassiz, 1850), Boreal Chorus Frog (IV)

Pseudacris triseriata (Wied-Neuwied, 1838), Western Chorus Frog (I, II, III, IV)

Family Ranidae (I, II, III, IV)

Rana catesbeiana Shaw, 1802, American Bullfrog (I, II, III, IV)

Rana clamitans melanota Rafinesque, 1820, Northern Green Frog (I, II, III, IV)

Rana palustris LeConte, 1825, Pickerel Frog (I, II, III, IV)

Rana pipiens Schreber, 1782, Northern Leopard Frog (I, II, III, IV)

Rana septentrionalis Baird, 1854, Mink Frog (III, IV)

Rana sylvatica LeConte, 1825, Wood Frog (I, II, III, IV)

Reptiles (I, II, III, IV):

Testudines--Turtles (I, II, III, IV)

Family Chelydridae (I, II, III, IV)

Chelydra serpentina serpentina (Linnaeus, 1758), Eastern Snapping Turtle (I, II, III, IV)

Family Emydidae (I, II, III, IV)

Chrysemys picta belli (Gray, 1831), Western Painted Turtle (IV)

Chrysemys picta marginata (Agassiz, 1857), Midland Painted Turtle (I, II)

Chrysemys picta belli x Chrysemys picta maginata (III, IV)

Clemmys guttata (Schneider, 1792), Spotted Turtle (I, II)

Emydoidea blandingii (Holbrook, 1838), Blanding's Turtle (I, II, III, IV)

Glyptemys insculpta (LeConte, 1830), Wood Turtle (I, II, III, IV)

Graptemys geographica (LeSueur, 1817), Northern Map Turtle (I, II, III, IV)

Terrapene carolina carolina (Linnaeus, 1758), Eastern Box Turtle (I, II)

Trachemys scripta elegans (Wied-Neuwied), Red-eared Slider (I, II)

Family Kinosternidae (I, II)

Sternotherus odoratus (Latreille, 1801), Stinkpot (formerly "Common Musk Turtle," see Crother et al. 2000) (I, II)

Family Trionychidae (I, II)

Apalone spinifera spinifera (LeSueur, 1827), Eastern Spiny Softshell (I, II)

Squamata--Lizards and Snakes (I, II, III, IV)

Family Scincidae (I, II, III, IV)

Eumeces fasciatus (Linnaeus, 1758), Common Five-lined Skink (I, II, III, IV)

Family Teiidae (I)

Aspidoscelis sexlineata (Linnaeus, 1766), Six-lined Racerunner (I)

Family Colubridae (I, II, III, IV)

Clonophis kirtlandii (Kennicott, 1856), Kirtland's Snake (I)

Coluber constrictor foxii (Baird and Girard, 1853), Blue Racer (I, II, III)

Diadophis punctatus edwardsii (Merrem, 1820), Northern Ring-necked Snake (I, II, III, IV)

Elaphe alleghaniensis (Holbrook, 1836), Eastern Rat Snakes (I, II)

Elaphe gloydi Conant, 1940, Eastern Foxsnake (I, II)

Elaphe vulpina (Baird and Girard, 1853), Western Foxsnake (III, IV)

Heterodon platirhinos Latreille, 1801, Eastern Hog-nosed Snake (I, II, IV)

Lampropeltis triangulum triangulum (Lacepede, 1788), Eastern Milksnake (I, II, III, IV)

Nerodia erythrogaster neglecta (Conant, 1949), Copper-bellied Watersnake (I)

Nerodia sipedon sipedon (Linnaeus, 1758), Common Watersnake (I, II, III, IV)

Opheodrys vernalis (Harlan, 1827), Smooth Greensnake (I, II, III, IV)

Regina septemvittata (Say, 1825), Queen Snake (I, II)

Storeria dekayi dekayi (Holbrook, 1836), Northern Brownsnake X Storeria dekayi wrightorum Trapido, 1944, Midland Brownsnake (I, II)

Storeria dekayi wrightorum Trapido, 1944, Midland Brownsnake (III, IV)

Storeria occipitomaculata (Storer, 1839), Northern Red-bellied Snake (I, II, III, IV)

Thamnophis butleri (Cope, 1889), Butler's Gartersnake (I, II)

Thamnophis sauritus septentrionalis (Linnaeus, 1766), Northern Ribbonsnake (I, II, IV)

Thamnophis sirtalis sirtalis (Linnaeus, 1758), Eastern Gartersnake (I, II, III, IV)

Family Viperidae (I, II)

Sistrurus catenatus catenatus (Rafinesque, 1818), Eastern Massasauga (I, II)

Differences in MRLE Amphibian and Reptile Species Diversity

Region I is, not unexpectedly, the most herpetologically diverse of all four regions in Michigan, having a total of 22 species of amphibians and 29 of reptiles (=51). Region II is next most diverse with a total of 19 species of amphibians and 27 of reptiles (=46). Region III has 18 species of amphibians and 15 of reptiles (=33), and Region IV has 18 species of amphibians and 16 of reptiles (=34).

Region I has 94.4% of the number of species that occur in the entire state of Michigan. The three species missing in Region I are Pseudacris maculata (occurs only on Isle Royale) and Rana septentrionalis, and Elaphe vulpina that occur only in the Upper Peninsula. Region II has 90.2% of the number of amphibian and reptiles species that occur in Region I. Region III has 64.7% and Region IV has 66.7% of the number of amphibian and reptile species that occur in Region I. Among other factors, these patterns probably reflect warmer temperatures throughout the year and greater habitat diversity in Region I than in the other three.

Comparing the regional diversity of amphibian species, we find that Region II has 86.4%, Region III 81.8%, and Region IV 81.8% of the number found in Region I. Comparing the regional diversity of reptile species, Region II has 93.1%, Region III only 51.7.1%, and Region IV only 55.2% of the number of species of reptiles found in Region I. In other words, there are only about half as many reptile species in Regions III and IV (Upper Peninsula) as there are in Region I. This may mainly reflect cooler temperatures and fewer habitats in Regions III and IV but there are probably other factors as well.

Post-glacial Re-entrant Taxa and the MRLE

About 20,000 yr. B.P. (years before present), the massive Wisconsinan Laurentide Ice Sheet moved southward to cover all of Michigan and extended southward to almost identical latitudes in southern Illinois, Indiana, and Ohio (Holman 1992, Figure 1, 454). This event completely obliterated all of the herpetological habitats in Michigan. Thus, all of the amphibians and reptiles that presently occur in Michigan have re-occupied the state since the ice began its final withdrawal about 14,800 yr. B.P.

Major physical factors associated with this herpetological re-occupation of Michigan include (1) the timing of the final deglaciation, (2) a warming trend that lasted from about 10,000 yr. B.P. to about 5,000 yr. B.P. and that peaked from about 7,000 to 5,000 yr. B.P. in an interval called the Hypsithermal (a warm, dry period), (3) a cooling trend that began about 5,000 yr. B.P. that peaked about 300 years ago in a short interval called the Little Ice Age (see Holman et al. 2003), and (4) the present warming trend that has occurred in Michigan from about 150 yr. B.P. to the present (Bernabo 1981).

Holman (1992) suggested two major routes of entry for the postglacial reoccupation of Michigan by amphibians and reptiles: (1) a wide corridor through Indiana and Ohio into the Lower Peninsula of Michigan; and (2) a somewhat narrower corridor through Wisconsin into the Upper Peninsula. Lesser invasions could have moved from Ontario into either one or both peninsulas, but it was suggested that few, if any, species were able to invade the Upper Peninsula from the Lower Peninsula across the Straits of Mackinac.

Secondly, Holman (1992) postulated the pattern and timing of the herpetological re-occupation of postglacial Michigan on the basis of the geological record, paleobotanical record, and modern and fossil ranges of species that presently occur in the state. Three categories of re-invading amphibians and reptiles were proposed. These (Table 1) are presented below with slight modifications of Holman (1992): Primary Invaders consisting of species that may be found in coniferous forest areas today, or, in one case (Rana sylvatica), in tundra; Secondary Invaders consisting of species that extend into the mixed conifer/broadleaf forest areas of the Lower Peninsula today but do not occur in the Upper Peninsula except for certain rare occurrences; and Tertiary Invaders consisting of those species that are all confined to the broadleaf area.

A comparison of the hypothetical Primary, Secondary, and Tertiary post-glacial re-invading herpetological species (modified somewhat from Holman [1992] based on new records) with the modern herpetofauna in the four Regions is as follows. Region I is unique in having Tertiary species that do not occur in any other region in the state. These Tertiary species are: Ambystoma opacum, A. texanum, Siren intermedia, Aspidoscelis sexlineata, Clonophis kirtlandii, and Nerodia erythrogaster. It is probable that these late re-invading species extended farther north during the mid-Holocene warm, dry spell (hypsithermal) and withdrew to their present ranges during the late Holocene cool spell that terminated in the Little Ice Age. Region II, on the other hand has only Primary and Secondary species.

Regions III and IV mainly contain Primary category amphibian and reptiles. These species are Ambystoma laterale, A. maculatum, A. tigrinum, Hemidactylium scutatum, Plethodon cinereus, Necturus maculosus, Notophtalmus viridescens, Bufo americanus, Hyla chrysoscelis, H. versicolor, Pseudacris crucifer, P. maculata, P. triseriata, Rana catesbeiana, R. clamitans, R. palustris, R. pipiens, R. septentrionalis, R. sylvatica, Chelydra serpentina, Chrysemys picta, Emydoidea blandingii, Glyptemys insculpta, Graptemys geographica, Eumeces fasciatus, Diadophis punctatus, Elaphe vulpina, Lampropeltis triangulum, Nerodia sipedon, Opheodrys vernalis, Storeria dekay, S. occipitomaculata, Thamnophis sauritus, and T. sirtalis.

Gaps in the Distribution of Widespread Herpetological Taxa

Region I. Beginning with the salamanders, a striking gap in distribution of three species of the burrowing salamander genus Ambystoma occurs. Ambystoma laterale is significantly absent among the dominant grain-producing counties in Michigan in Region I. These include Isabella, Midland, Bay, Saginaw, and Tuscola in the Midland and Saginaw Bay area; and in Lenawee and Monroe counties in extreme southeastern Michigan (see Sommers 1977, 148). These counties not only are among the most intensely agriculturalized counties in the state but are also grouped among the least forested counties in the state (see Dickmann and Leefers 2003, Figure 1.2., 6). Another factor contributing to the absence of this species in the Midland-Saginaw Bay area group of counties may be the acidic fossil dune and dune soils in these areas (Holman 2001b). Geologically, all of these counties occur in glacial lake plains (Dorr and Eschman 1970).

Probably reflecting the same factors above are Ambystoma maculatum, which is absent from the same five counties in the Midland-Saginaw Bay area and from Monroe County in extreme southeastern Michigan, and Ambystoma tigrinum, found only in the bottom three tiers of counties in Region I (except for St. Joseph, Lenawee, Monroe, and Macomb counties in southeastern Michigan).

Turning to the lungless salamander genera Hemidactylium and Plethodon, each represented by a single species in Michigan, we find that Hemidactylium scutatum is missing from the previously discussed counties in the Midland area as well as all of the counties surrounding Saginaw Bay. Moreover, H. scutatum has not been recorded from Allegan, Van Buren, Kalamazoo, Berrien, and Cass counties in southwestern Michigan and extending across the entire lowest tier of counties in the state. Plethodon cinereus, the most abundant salamander in the state, occurs in every county in Michigan except for Saginaw, Tuscola, and Sanilac.

Relative to the anurans, the presence of sandy soils, and possibly temperature gradients, appears to be a factor in the distribution of Bufo fowleri. In Region I this species occurs in roughly the relatively warm and sandy western half of the state and is absent in the eastern half of the state except for Monroe and Wayne counties. Acris crepitans, presently occurring only in Region I (with the exception of a single extralimital record from Leelanau County in Region II) is absent from most of the counties in the Saginaw Bay area and the thumb of Michigan, including Bay, Huron, Tuscola, Genesee, Sanilac, and St. Clair.

Sternotherus odoratus is widespread in Region I, but conspicuously absent in the Saginaw Bay-Thumb counties of Bay, Huron, Tuscola, Sanilac, Lapeer, and St. Clair. It has also not been recorded in the extreme southeastern counties of Wayne and Monroe. I suspect it is because of a general lack of lakes with shallows of marl, sand, or gravel bottoms (see Harding and Holman 1990) in these areas. Apalone spinifera has not been recorded from any modern sites in the Midland-Sagninaw Bay-Thumb areas including Midland, Bay, Saginaw, Tuscola, Huron, Sanilac, Lapeer, and St. Clair.

Relative to the lizards, although Eumeces fasciatus occurs in all four regions in the state, its distribution is particularly spotty in Region I. There is a large gap in the occurrence of this lizard in the western portion of Region I in counties that do not border Lake Michigan, and it has been recorded only in Huron and St. Clair counties in the thumb.

Turning to snakes, Coluber constrictor is conspicuously absent in the Midland-Saginaw Bay-Thumb area counties, including Midland, Bay, Saginaw, Tuscola, Huron, Sanilac, Lapeer, and St. Clair. Records of Diadophis punctatus are relatively rare in Region I, being confined to four counties in the western part of the state (Muskegon, Kent, Allegan, and Cass), and three counties in the southeastern part of the region (Jackson, Washtenaw, and Oakland). Elaphe alleghaniensis occurs in all of the counties bordering Lake Michigan in the west, but is absent from the Midland-Saginaw Bay-Thumb areas except for Lapeer County, and is of somewhat spotty occurrence everywhere else in Region I. Heterodon platirhinos is represented well in much of the western portion of Region I and in the southeastern portion (Oakland, Washtenaw, Wayne, and Monroe counties) but has not been recorded in most of the fifth tier of counties and all of the thumb region, except for Huron County. It is of interest that this species has been found in Monroe County, one of the most highly agriculturalized counties in the state.

Nerodia sipedon has not been recorded in the Saginaw Bay-Thumb area counties of Bay, Tuscola, Sanilac, and St. Clair. Opheodrys vernalis occurs throughout Michigan but is relatively uncommon in Region I. In fact, I have not seen it in this region for 36 years. The only concentrated group of county records in the area is in Jackson, Washtenaw, Livingston, Oakland, and St. Clair counties in the southeastern part of the region. Regina septemvittata has not been recorded in the upper central counties of Region I as well as most of the Midland-Saginaw-Bay-Thumb area and the central part of the lower two tiers of counties. Storeria occipitomaculata is uncommon in the western part of Region I, occurring only in Allegan County. On the other hand, it is relatively common in the eastern part of Region I, including the Midland-Saginaw Bay-Thumb except for Huron and Sanilac counties. No records exist for the lowest two tiers of counties except for Washtenaw and Wayne counties. Interestingly, a very similar distributional situation occurs in Thamnophis butleri. Michigan's only venomous snake, Sistrurus catenatus, has been recorded widely in Region I except for most of the Midland-Saginaw Bay-Thumb area, where it has only been recorded from Huron County.

Region II. Here we find a strange gap in the range of Ambystoma maculatum which has not been recorded from the second and third tiers of counties from the bottom of the region except for Mason and Manistee that border Lake Michigan.

Turning now to the lizards, Eumeces fasciatus is of much more widespread occurrence in Region II than Region I, but has not been recorded from four counties in northwestern Region II that border Lake Michigan. These are Leelanau, Antrim, Charlevoix, and Emmet.

Relative to the snakes, Coluber constrictor follows the Traverse Corridor (a warm belt along coastal Michigan) north to Benzie and Grand Traverse counties. Two other records in Region II, Clare County in the lower central part of the region and Otsego County near the tip of the Lower Peninsula, are disjunct. Storeria occipitomaculata has not been recorded from Oceana, Newaygo, Mecosta Lake, Manistee, Wexford, Benzie, and Leelanau counties, but is present in all of the other counties in Region II. Thamnophis sauritus is relatively widespread in Region II, with the exception of a lack or records in three counties, Arenac, Iosco, and Alcona, that border Lake Huron in the eastern part of the region.

Region III. The turtle Emydoidea blandingii is absent in the eastern three counties, Chippewa, Mackinac, and Luce, but present in the remaining counties of the region. The lizard Eumeces fasciatus follows the same odd pattern, being absent in the eastern four counties of Region III (Chippewa, Mackinaw, Luce, and Schoolcraft), but present in the western Region III counties of Alger and Delta. The snake Elaphe vulpina follows this same pattern, being absent in the eastern three counties of Region III and present in the remaining ones.

Region IV. Emydoidea blandingii has not been recorded in five western counties, including Baraga, Gogebic, Ontonagon, Houghton, and Keweenaw. Eumeces fasciatus is also absent from all but the eastern part of Region IV, occurring only in Marquette, Dickinson, and Menominee counties. Relative to the snakes, a single record of Heterodon platirhinos occurs in Region IV in southwestern Menominee County, but this record is contiguous with populations of this snake in Wisconsin. Nerodia sipedon has a large gap in its range in Region IV as it occurs only in the westernmost two counties in the region (Gogebic and Ontonagon) and in Menominee County in the eastern part of Region IV.

Relict Species

I follow here the definition of a relict species proposed by Lincoln et al. (1982, p. 216), namely that relict species are "Persistent remnants of formerly widespread fauna or flora existing in isolated areas or habitats."

Region I. Turning first to the salamanders, Ambystoma opacum is part of an isolated population that is confined to Berrien, Van Buren, and Allegan counties in extreme southwestern Michigan and the Lake Michigan dunes of Porter and LaPorte counties in extreme northwestern Indiana (Minton 2001). All five counties are contiguous and border Lake Michigan. Farther south in Indiana the species is widespread. I believe the most reasonable hypothesis is that this population is relict from the warmer hypsithermal interval of the Holocene and has been able to maintain itself do to the warming effect of nearby Lake Michigan. Ambystoma texanum has been recorded in Michigan only in Region I in the far southeast corner of the state, namely Livingston, Washtenaw, Wayne, Hillsdale, and Monroe. This cannot be considered a relict population as it is contiguous with the widespread population of the species in Ohio (Pfingsten 1998, Figure 25-10, 228). It seems possible that the range of this species could have extended northward in Michigan during the hypsithermal interval. Siren intermedia nettingi is found only in Region I and has been recorded from only two counties, Van Buren and Allegan in southwestern Lower Michigan. Since this species is not known in the upper tier of counties in Indiana, except for a record in extreme southwestern Porter County (Minton 2001, map, 102), I suggest that S. intermedia is part of a relict population in Michigan that may have extended farther north in the state along the Traverse Corridor during the hypsithermal interval.

Relative to turtles, the modern range of Trachemys scripta elegans is from Texas east to Mississippi and western Alabama and north to western Indiana through western Tennessee and most of Kentucky. There are large isolated populations of T. s. elegans in southern Ohio and eastern Maryland (Conant and Collins 1998), and smaller isolated, possibly relict ones in Regions I and II in Michigan (Holman 1994). This subspecies of turtle is unique in that released specimens from the pet trade occur throughout the world in temperate and tropical areas. One could attribute these populations in Michigan to such releases, except there are arguments that some of these populations may be relicts from hypsithermal times (Holman 1994). A record of this species from an archaeological site in Saginaw County (Shultz Site, ca 2,500-1600 years ago) was recovered from among the refuse of local animals used as food. Adler (1968) suggested that this record and another record from the Durst Archaeological Site at about the same latitude in southcentral Wisconsin indicated that the range of the Red-eared Slider has been reduced in subsequent times.

Concerning the lizards, Aspidoscelis sexlineata is known in Michigan only on the basis of a single but thriving population in Tuscola County. Holman (2000b) suggested that the population is a relict one left over from warm, dry (hypsithermal) times in the mid-Holocene. In about 1800 a corridor composed of tree punctuated grasslands (grasslands interspersed by open areas of gnarled oak species) extended from eastern Berrien County in the southeastern most corner of Michigan across the southern part of the state to within 20 kilometers of Tuscola County (Dickmann and Leefers 2003, map p. 102 and pp. 103-4). This 1800 Michigan corridor would connect today with the dune and prairie areas in northwestern Indiana in Lake, Porter, Laporte, Newton, Jasper, Pulaski, and Starke counties, all of which presently support populations of Aspidoscelis sexlineata (Minton 2001). Certainly the warmer, dryer hypsithermal interval in the mid-Holocene would have made this "corridor" even more accessible to this prairie species.

The snake Clonophis kirtlandii is found in Michigan only in Region I where records are known in southwestern Michigan in Berrien, Cass, Van Buren, Kalamazoo, Ottawa, and southern Muskegon County; and in Lenawee and Washtenaw counties in extreme southeastern Michigan. These are not relict populations, at least in the broad sense, but northern extensions of adjacent scattered populations in Indiana and Ohio (see Minton 2001, map p. 279 and Conant 1951, map 17, 75). It is not impossible that this small snake extended farther north during hypsithermal times in the state. Nerodia erythrogaster neglecta occurs in southern Michigan only in Cass, St. Joseph, Kalamazoo, Branch, Hillsdale, Eaton, and Oakland counties as a part of a large relict population that is contiguous in northeastern Indiana and northwestern Ohio (see Conant and Collins 1998; Minton 2001; and Conant 1951).

Region II. Starting with salamanders, there are only isolated finds of Ambystoma tigrinum in the area. This species occurs in two counties, Crawford and Otsego, in the northcentral part of the region, Manistee county adjacent to Lake Michigan, and Clare County in the southcentral part of the region. These are probably small relict populations, as only the bottom three tiers of counties in Region I have records of this species. Moreover, all of Regions III and IV (other than an odd mainly larval population in III) lack records for this species.

Relict populations of Graptemys geographica occur in Roscommon and Grand Traverse counties, as no other records of this species are known north or directly south of these counties in Region II. A very isolated relict population of Sternotherus odoratus occurs in the northeastern part of the region, as the species is known from only a single locality in Montmorency County. The nearest county record of the above species in Michigan is in Newaygo County, far to the southwest. The same situation exists for Apalone spinifera which is known from Grand Traverse and Crawford counties in the northern part of the area. Again, the nearest modern record of the species is from Newaygo County to the southwest.

A clearly relict population of the snake Regina septemvittata occurs on Bois Blanc Island, in Lake Huron near Cheyboygan in the extreme northern part of the area. The nearest populations are in Manistee County to the southwest and Ogemaw County to the southeast. Thamnophis butleri exists in what is probably a relict population in Presque Isle and Alpena counties in the northeastern portion of the area. The nearest record is Arenac county to the south.

Region III. Relative to salamanders, a single, mainly larval population of Ambystoma tigrinum from Alger County represents the only record of this species in the Upper Peninsula. Incidentally, one of these specimens (Hensley 1964) was 240 mm long! The nearest locality for the species is from Otsego County in Region II, approximately 282 kilometers south and east of the Alger County locality (Hensley 1964).

A probable relict population of the turtle Graptemys geographica is known from east-central Schoolcraft County. The nearest known population of this species (also a probable relict population) is known from Dickinson County to the west.

Turning to relict snake populations in Region III, a population of Coluber constictor occurs in Menominee Country. This is obviously a remarkable relict population, as otherwise this species is confined to southwestern Wisconsin to the southwest (Vogt 1981) and is well separated from this species in the eastern part of Region II. Lampropeltis triangulum is known in the region only in the eastern part of Mackinac County; this population should be considered a relict one until additional collecting can prove otherwise. Storeria dekayi wrightorum occurs in Schoolcraft and Delta counties in the region. The nearest occurrence to these populations is in southwestern Marinette County, Wisconsin (Vogt 1981).

Region IV. The turtle Graptemys geographica is known in the region only in Dickinson County. This is a probable relict population as it is not found in any of the three tiers of Wisconsin counties to the southwest (see Vogt 1981, map, 106).

A relict population of the snake Lampropeltis triangulum occurs in the upper tip of Marquette County, far from any other records in Michigan or Wisconsin. A relict population of Storeria dekayi wrightorum occurs in the isolated area of Baraga and Keewenaw counties in the northwestern part of the region.

SUMMARY

The four Michigan Regional Landscape Ecosystems are based mainly on glacially derived landforms, the underlying bedrock, soils, and climates. The herpetofauna of each of these regions was determined based on museum specimens and records from ongoing herpetological studies. These regional herpetofaunas were compared on the basis of taxonomic diversity; the presence of Primary, Secondary, and Tertiary re-entrant post-glacial species of Holman (1992); the presence of gaps in the distribution of generally widespread taxa; and the presence of relict species.

Region I has the most taxonomically diverse herpetofauna with 22 species of amphibians and 29 reptiles (total 51). Region I dominated the other regions in having six Tertiary species. It also dominated the other regions in having 20 species with significant gaps in their distribution (39.2% of its species). Region I has five relict species.

Region II has the next most taxonomically diverse herpetofauna with 19 species of amphibians and 27 reptiles for a total of 46 (90.2% of those in Region I). Region II has only Primary and Secondary species and only five species with significant gaps in their distribution. Region II has six relict species, one more than in Region I.

Regions III and IV are more similar to one another both taxonomically and in lack of species diversity (especially in reptiles) than Region I compared to Region II. The herpetological similarity of III and IV is of considerable interest considering that the two regions have important climatic, topographic, geologic, and soil differences.

Region III has 18 species of amphibians and 15 reptiles (total of 33) and IV has 18 species of amphibians and 16 reptiles (total of 34)--this is 64.7% and 66.7% of the total species found in Region I. Only Primary amphibians and reptiles occur in III and IV with the exception of two Secondary reptiles in Region III. Regions III and IV each have three widespread species with gaps in their distribution. Region III has five relict species and Region IV has three.

APPENDIX

County records of Michigan amphibian and reptile species (in sequence). An "A" rather than a shaded area indicates that the record in that county is from an archaeological site only.

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TABLE 1. Primary (P), Secondary (S), and Tertiary (T) amphibians and
reptile species re-occupying Post-Glacial Michigan (modified from
Holman, 1992). A = All Regions. Individual MRLE occurrences symbolized
by Roman numerals.

Ambystoma laterale P, Chelydra serpentina P, Regina septemvittata S,
A A I, II
Ambystoma maculatum Chrysemys picta P, A Storeria dekayi P, A
P, A
Ambystoma opacum T, I Clemmys guttata S, I, Storeria occipitomaculata
 II P, A
Ambystoma texanum T, Emydoidea blandingii Thamnophis butleri S, I,
I P, A II
Ambystoma tigrinum P, Glyptemys insculpta P, Thamnophis sauritus P, I,
I, II, III A II, IV
Hemidactylium Graptemys geographica Thamnophis sirtalis P, A
scutatum P, A P, A
Plethodon cinereus P, Terrapene carolina S, Sistrurus catenatus S, I,
A I, II II
Necturus maculosus P, Trachemys scripta S,
A I, II
Notopthalmus Sternotherus odoratus
viridescens P, A S, I, II
Siren intermedia T, I Apalone spinifera S,
 I, II
Bufo americanus P, A Eumeces fasciatus P, A
Bufo fowleri S, I, II Aspidoscelis
 sexlineata T, I
Acris crepitans S, I,
II
Hyla chrysoscelis P, Clonophis kirtlandii
A T, I
Hyla versicolor P, A Coluber constrictor S,
 I, II, III
Pseudacris crucifer Diadophis punctatus P,
P, A A
Pseudacris maculata Elaphe alleghaniensis
P, IV S, I, II
Pseudacris triseriata Elaphe gloydi S, I, II
P, A
Rana catesbeiana P, A Elaphe vulpina P, III,
 IV
Rana clamitans P, A Heterodon platirhinos
 S, I, II, III
Rana palustris P, A Lampropeltis
 triangulim P, A
Rana pipiens P, A Nerodia erythrogaster
 T, I
Rana septentrionalis Nerodia sipedon P, A
P, III, IV
Rana sylvatica P, A Opheodrys vernalis P,
 A


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J. ALAN HOLMAN

Michigan State University
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