SPHENOPHRYNE - American Museum of Natural History

SPHENOPHRYNE - American Museum of Natural History SPHENOPHRYNE - American Museum of Natural History

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6 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 253 to transforming my crude sketches into realistic drawings. METHODS ANATOMY: On most specimens examined I made a suite of measurements with dial calipers read to the closest 0.1 mm or, if appropriate, I used an ocular micrometer in a binocular dissecting microscope read to the nearest 0.05 mm. Sex of the specimen was apparent if the specimen was a male calling when captured. Otherwise, males were determined by the presence of vocal sac openings (absent in two species) or by examination of gonads, females were determined by examination of gonads. With the exception of a few dried preparations, descriptions of osteology derive from cleared and stained specimens (identified as ‘‘C&S’’ in lists of specimens examined). Color descriptions of living frogs are from my field notes and photographs, supplemented by notes and photographs provided by other collectors. The following abbreviations pertain to measurements made (with some exceptions) on each specimen: EN. Distance between anterior edge of eye opening and center of external naris. EY. Distance between anterior and posterior edges of eye opening. It is sometimes necessary to push the eyeball up from within the mouth in order to approximate the condition in life. FD. Width of disc of third finger measured at a right angle to the axis of the digit with the disc flattened against a glass plate. FT. Length of foot between proximal edge of inner metatarsal elevation and tip of fourth toe (see HD). HD. Length of hand between proximal edge of inner metacarpal elevation and tip of third finger. Both hand and foot measurements may have reduced accuracy owing to the sometimes indistinct nature of the metacarpal and metatarsal elevations and to the difficulty of properly spreading the hands and feet of indifferently preserved specimens. HW. Head width at widest point, generally at the level of the tympanum or jaw angle. IN. Distance between centers (not medial edges) of external nares. SVL. Length from snout to vent—from tip of snout to cloacal opening, with body flattened if necessary. TD. Width of disc of fourth toe measured at a right angle to the axis of the digit with the disc flattened against a glass plate. TL. Tibia length, between heel and outer surface of flexed knee. TY. Diameter of tympanum, including tympanic ring, measured horizontally. In most species the ear is scarcely apparent externally and hence it is difficult to measure with accuracy. Relative lengths of fingers and toes were determined by placing them parallel to the third finger or fourth toe; actual lengths were not measured. USE OF PROPORTIONS Species of frogs usually differ in one or more body proportions, and these differences may be employed usefully in discriminating among species. Differences may be evident in comparisons of regressions of the measurements of body parts against body size (SVL), although overlap in ranges may hinder the use of regression data in species discrimination. I routinely calculated and plotted regression data for samples of all species. Ratios comparing body part size to SVL, of course, suffer from the same difficulty; that is, overlap between species ranges of ratios may be so great that the differences in proportions, although statistically and possibly biologically significant, may be largely useless for specific identification of individuals. An effective method of overcoming this problem utilizes two different ratios simultaneously, in each of which the two species being compared show different but overlapping ranges of variation. The statement takes this form or a variant thereof: ‘‘Ratio A less than X and ratio B greater than Y, vs. ratio A greater than X and ratio B less than Y.’’ A graphic presentation allows one to place a given specimen in the spectrum of points. Identifications may be greatly facilitated in this manner. Ratios present another problem, however. Most proportions—at least those commonly

2000 ZWEIFEL: PARTITION OF SPHENOPHRYNE 7 used in systematics studies—change with growth. The degree and direction of change varies not only interspecifically, but also with the body part being considered. By using ratios of adults only, the ranges of variation can often be considerably reduced, leading to less overlap among species and more utility in keying. Ontogenetic change in proportions is discussed under Morphology. LOCALITY RECORDS A variety of circumstances hamper, even frustrate, fixing of localities of collection in New Guinea. Even in comparatively recently it has been difficult for field workers remote from towns or the few roads to pinpoint their location. Even as comprehensive maps become available (especially for Papua New Guinea), and with the increasing use of Geographic Positioning System technology, there remains a legacy of politically based and other sources of confusion in museum catalogs and published papers. IRIAN JAYA: From the 1800s until 1963, the western half of New Guinea was under Dutch control; ‘‘Netherlands New Guinea’’ and ‘‘Dutch New Guinea’’ were the commonly used names. Following the Indonesian assumption of control, the name ‘‘Irian Barat’’ saw some usage (at least as late as 1967). This was followed by ‘‘Irian Jaya,’’ as a Province of Indonesia. Quite recently the name was changed again to ‘‘Papua Barat’’ (or West Papua). In the present work I continue to use the more familiar Irian Jaya. Administrative subdivisions of Irian Jaya exist but rarely appear in museum records. Many geographic names established under Dutch governance have been replaced by Indonesian ones. For example, the principal city was Hollandia, then Sukarnapura, then Jayapura, now Port Numbai. PAPUA NEW GUINEA: The eastern half of New Guinea has had a more complicated political history as well as much more biological collecting activity than has Irian Jaya. The first administrative organization of the northern part was by Germany in 1884 as Kaiser Wilhelmsland. This also included islands to the north and east—Neu Pommern (New Britain), Neu Mecklenburg (New Ireland), and many smaller islands. German New Guinea and Northeast New Guinea are other names often encountered in the literature and in museum records. Australia occupied Kaiser Wilhelmsland in 1914, and in 1920 the League of Nations mandated administrative control of the area to Australia as the Trust Territory of New Guinea. The southern part of Papua New Guinea was originally (1888) British New Guinea; in 1906 it was turned over to Australian administration and renamed Papua, later called the Territory of Papua but governed separately from the Territory of New Guinea. Governmental evolution subsequent to World War II produced the combined Territory of Papua New Guinea and then (1975) the independent nation of Papua New Guinea. The territories of Papua and New Guinea were subdivided into administrative districts that, with nationhood, were redesignated as provinces. Many of these did not long retain their original integrity. New provinces were carved out of old, boundaries were adjusted, provinces were renamed, and alternate spellings were used. Even names used by the provincial and central governments disagree in some instances. For Papua New Guinea, I have standardized province records according to the names

6 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 253<br />

to transforming my crude sketches into realistic<br />

drawings.<br />

METHODS<br />

ANATOMY: On most specimens examined I<br />

made a suite <strong>of</strong> measurements with dial calipers<br />

read to the closest 0.1 mm or, if appropriate,<br />

I used an ocular micrometer in a binocular<br />

dissecting microscope read to the<br />

nearest 0.05 mm. Sex <strong>of</strong> the specimen was<br />

apparent if the specimen was a male calling<br />

when captured. Otherwise, males were determined<br />

by the presence <strong>of</strong> vocal sac openings<br />

(absent in two species) or by examination<br />

<strong>of</strong> gonads, females were determined by<br />

examination <strong>of</strong> gonads. With the exception<br />

<strong>of</strong> a few dried preparations, descriptions <strong>of</strong><br />

osteology derive from cleared and stained<br />

specimens (identified as ‘‘C&S’’ in lists <strong>of</strong><br />

specimens examined). Color descriptions <strong>of</strong><br />

living frogs are from my field notes and photographs,<br />

supplemented by notes and photographs<br />

provided by other collectors.<br />

The following abbreviations pertain to<br />

measurements made (with some exceptions)<br />

on each specimen:<br />

EN. Distance between anterior edge <strong>of</strong><br />

eye opening and center <strong>of</strong> external naris.<br />

EY. Distance between anterior and posterior<br />

edges <strong>of</strong> eye opening. It is sometimes<br />

necessary to push the eyeball up from within<br />

the mouth in order to approximate the condition<br />

in life.<br />

FD. Width <strong>of</strong> disc <strong>of</strong> third finger measured<br />

at a right angle to the axis <strong>of</strong> the digit<br />

with the disc flattened against a glass plate.<br />

FT. Length <strong>of</strong> foot between proximal edge<br />

<strong>of</strong> inner metatarsal elevation and tip <strong>of</strong> fourth<br />

toe (see HD).<br />

HD. Length <strong>of</strong> hand between proximal<br />

edge <strong>of</strong> inner metacarpal elevation and tip <strong>of</strong><br />

third finger. Both hand and foot measurements<br />

may have reduced accuracy owing to<br />

the sometimes indistinct nature <strong>of</strong> the metacarpal<br />

and metatarsal elevations and to the<br />

difficulty <strong>of</strong> properly spreading the hands<br />

and feet <strong>of</strong> indifferently preserved specimens.<br />

HW. Head width at widest point, generally<br />

at the level <strong>of</strong> the tympanum or jaw angle.<br />

IN. Distance between centers (not medial<br />

edges) <strong>of</strong> external nares.<br />

SVL. Length from snout to vent—from<br />

tip <strong>of</strong> snout to cloacal opening, with body<br />

flattened if necessary.<br />

TD. Width <strong>of</strong> disc <strong>of</strong> fourth toe measured<br />

at a right angle to the axis <strong>of</strong> the digit with<br />

the disc flattened against a glass plate.<br />

TL. Tibia length, between heel and outer<br />

surface <strong>of</strong> flexed knee.<br />

TY. Diameter <strong>of</strong> tympanum, including<br />

tympanic ring, measured horizontally. In<br />

most species the ear is scarcely apparent externally<br />

and hence it is difficult to measure<br />

with accuracy.<br />

Relative lengths <strong>of</strong> fingers and toes were<br />

determined by placing them parallel to the<br />

third finger or fourth toe; actual lengths were<br />

not measured.<br />

USE OF PROPORTIONS<br />

Species <strong>of</strong> frogs usually differ in one or<br />

more body proportions, and these differences<br />

may be employed usefully in discriminating<br />

among species. Differences may be evident<br />

in comparisons <strong>of</strong> regressions <strong>of</strong> the measurements<br />

<strong>of</strong> body parts against body size<br />

(SVL), although overlap in ranges may hinder<br />

the use <strong>of</strong> regression data in species discrimination.<br />

I routinely calculated and plotted<br />

regression data for samples <strong>of</strong> all species.<br />

Ratios comparing body part size to SVL, <strong>of</strong><br />

course, suffer from the same difficulty; that<br />

is, overlap between species ranges <strong>of</strong> ratios<br />

may be so great that the differences in proportions,<br />

although statistically and possibly<br />

biologically significant, may be largely useless<br />

for specific identification <strong>of</strong> individuals.<br />

An effective method <strong>of</strong> overcoming this<br />

problem utilizes two different ratios simultaneously,<br />

in each <strong>of</strong> which the two species<br />

being compared show different but overlapping<br />

ranges <strong>of</strong> variation. The statement takes<br />

this form or a variant there<strong>of</strong>: ‘‘Ratio A less<br />

than X and ratio B greater than Y, vs. ratio<br />

A greater than X and ratio B less than Y.’’<br />

A graphic presentation allows one to place a<br />

given specimen in the spectrum <strong>of</strong> points.<br />

Identifications may be greatly facilitated in<br />

this manner.<br />

Ratios present another problem, however.<br />

Most proportions—at least those commonly

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