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A t american museum

Novitates

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3628, 19 pp., 46 figures, 1 table September 10, 2008

Classification, Natural History, and Evolution
of Epiphloeinae (Coleoptera: Cleridae). Part V.
Decorosa Opitz, a New Genus of Checkered Beetles
from Hispaniola with Description of Its Four

New Species

WESTON OPITZ 1

ABSTRACT

The new genus Decorosa Opitz, known only from the Dominican Republic, and its four new
species are described: D. aladecoris, D. iviei, D. limatula, and D. neiba. Reduction of the seventh
row of elytral punctations and filiform funicular antennomeres distinguish Decorosa specimens
from other epiphloeines with antennae composed of 10 antennomeres. Aside from the
conventional components of generic revisions this treatise also includes a review of Greater
Antillean and Hispaniolan paleogeography, and comments about phylogenetics and zoogeo¬
graphic considerations.

Distributional records, in conjunction with analysis of intrageneric relationships suggest that
Decorosa evolved in northern Hispaniola. Available specimens of Decorosa were collected from
mountain forests of the Cordillera Central (1160-2885 m), Sierra de Neiba, (1856-1874 m), and
from eastern terrain prehistorically considered part of the northern element of ancient Hispaniola.
Compelling evidence from geology and biology suggests that extant Hispaniola was prehistorically
composed of a northern and southern island. It is suggested that ancestral Decorosa split into the
limatula-iviei and decorosa-neiba stocks in the more northern ancient terrain of Hispaniola.
Subsequently, each of two ancestral lineages yielded at least two sister species. It is further
postulated that there occurred two dispersal events that brought one species of each sister pair to
more southern Hispaniola; limatula from the limatula-iviei lineage and neiba from the aladecoris-
neiba lineage.

1 Kansas Wesleyan University, Department of Biology, 100 East Claflin Avenue, Salina, KS 67401-6196
(opitz@kwu.edu).

ISSN 0003-0082

AMERICAN MUSEUM NOVITATES

NO. 3628

RESUMEN

El genero nuevo Decorosa, de distribution restringida a Republica Dominicana, y sus cuatro
nuevas especies son descritos: D. aladecoris, D. iviei, D. limatula, y D. neiba. Tres caracteristicas
discriminan los especimenes de Decorosa de otros Epiphloeinae: antenas compuestas de 10
antenomeros, reduccion de la septima linea de puntuaciones y antenomeros funiculares filiformes.
Ademas de los elementos convencionales de una revision generica, este trabajo incluye una revision
de la paleogeografia de Hispaniola y las Grandes Antillas, asi como comentarios sobre la filogenia
y la zoogeografia del genero. Cuarenta dibujos, incluyendo cuatro de especimenes enteros, una
micrografia electronica, una fotografia, un mapa y un arbol filogenetico ilustran el trabajo. Las
caracteristicas morfologicas son polarizadas filogeneticamente, organizadas en una matriz de
caracteres y analizadas en una hipotesis filogenetica intragenerica, contrastada con un outgroup
compuesto por los demas generos de Epiphloeinae con antenas de 10 antenomeros.

La distribucion geografica, en conjunto con el analisis de relaciones intragenerico, sugiere que el
grupo ancestral de Decorosa incluye el norte de Hispaniola. Los especimenes disponibles de
Decorosa fueron colectados en las selvas de montanas de la Cordillera Central (1160-2885 m), en
la Sierra de Neiba (1856-1874 m), y en terrenos del este, considerados prehistoricamente como
parte del elemento norte de la antigua Hispaniola. Existe evidencias geologica y biologica que
sugieren que la actual Hispaniola era prehistoricamente compuesta de una isla norte y una isla sur.
Se sugiere que los ancestros de las especies del genero Decorosa se separaron en el grupo limatula-
iviei y el grupo decorosa-neiba en la parte mas norte de Hispaniola. Posteriormente, cada linaje dio
origen a por lo menos dos especies hermanas. Se postula que ocurrieron dos eventos de dispersion,
que empujaron una especie de cada par de especies hermanas mas hacia el sur de Hispaniola;
limatula del linaje limatula-iviei y neiba del linaje aladecoris-neiba.

INTRODUCTION

Despite the long history of entomological
research in Hispaniola there remains a dearth
of information about the diversity of Cleridae
on the island. Not one species is listed from
that island in the world catalogue of
Corporaal (1950). I became interested in the
entomofauna of this island as a result of my
studies of the new world subfamily Epiphloeinae,
which occur on many of the Caribbean islands.

My commitment to make knowledge avail¬
able incrementally about this monophyletic
lineage of the Cleridae, the epiphloeines, has
involved lengthy periods of specimen accumu¬
lation. Insufficient number of specimens may
at times be a considerable deterrent toward
credible hypotheses of the biological species. It
also minimizes the opportunity for discussions
of a taxon’s evolution and biogeography. This
is particularly relevant among the anatomi¬
cally homogeneous components of epiphloeine
species (Opitz, 2004: 7). Adequately represent¬
ing the full geographic range of these check¬
ered beetles is imperative for their biological
study so that a hypothesis about their species
concept may be credible and, thus, their
ecological role may be most usefully evaluat¬
ed. Extraordinary efforts in cooperation from
colleagues have greatly facilitated this goal.

Their loans, exchanges, and gifts have greatly
enhanced my research progress.

Elsewhere, I (Opitz, 2005: 8; 2007: 79) allude
to difficulties one usually encounters in
attempts to collect Cleridae, with frequent
capture events of a single individual or no
specimens at all. Therefore, loans from muse¬
ums and private collections have been the
primary source of my research material. In
recent months, I was delighted to receive an
assortment of checkered beetles from Robert
Davidson, Michael A. Ivie, and Philip Perkins,
on whose specimens this study is based.

MATERIALS AND METHODS

This contribution is based on 16 adult
specimens of Decorosa. Unfortunately, small
sample sizes are common in taxonomic
revisions of the Cleridae because species of
many genera are often collected in increments
of one. One specimen of D. iviei, n. sp., the
type species, was completely disarticulated to
establish the basic anatomical characteristics
of the genus. All male specimens were
dissected to determine aedeagal and spicular
fork structure. The primary type specimen of
Decorosa aladecoris, n. sp., was prepared with
legs and antennae symmetrically arranged for
photography with a Nikon DXM1200 digital

2008

OPITZ: EPIPHLOEINAE, PART V: DECOROSA OPITZ

camera attached to a Leica MZ microscope.
The scale bar with morphological illustrations
represent 1 millimeter.

The rationale to identify biological species
and to predict phylogenetic relationships is
similar to those expressed in earlier works
(Opitz, 2006: 84. 2007: 84). I use “homoplasic
apotypy” to define those characteristics that I
presume evolved independently in evolutionari-
ly distant taxa conglomerates. The rational and
justification for the use of such apotypies will be
addressed elsewhere (Opitz, in preparation).

Techniques of dissection, measurements,
illustration, and terminology of anatomic
structure are similar to those described in my
previous revisionary works (Ekis, 1977: 6;
Opitz, 2004: 7; 2005: 8). Label information of
each primary type is presented in species
descriptions in exact sequence and script as
indicated on specimen labels. In descriptions, I
refer to primary (1°) and secondary (2°) degree
setae not in the sense that they might be used in
larval ontogeny, but in the sense of physical
prominence, with primary setae being much
longer than the secondary setae. These setae are
found on the elytral disc and are further defined
as follows: primary setae are longer and more
suberect than secondary setae, which are short
to an extent that they give an impression of
“setal mats” (Opitz, 2005: 80, fig. 260). Rows of
punctations on the elytral disc are numbered
such that the row nearest the sutural margin is
considered row number 1. Species descriptions
also include an itemization of the kinds of
labels found beneath the holotype, further
minimizing potential confusion of the primary
type identity during future research. Specific
epithets were coined or taken from the com¬
prehensive work of Brown (1956).

Sister group relationships within Decorosa
are based on principles as advocated by
Hennig (1966), but I agree with Tuomikosky
(1967: 138) who states that the terms apotypic
and plesiotypic rather than apomorphic and
plesiomorphic should be used on the grounds
that evolutionary syntheses are not confined
to structural character states. The apotypic or
plesiotypic states of adult morphological
characters are determined and organized into
a character matrix. The data matrix was
analyzed by hand, which represents strictly
intuitive conclusions (fig. 44), and by use of

NONA (Goloboff, 1993) in conjunction with
Winclada version 1.00.08 (Nixon, 2002) (fig. 45).

Specimens are deposited in the following
collections and copies of this contribution will
be sent to their respective libraries:

AMNH American Museum of Natural

History, Department of Entomology,
Central Park West at 79th Street, New
York, New York 10024-5192 (Lee
Herman; herman@amnh.org).
CMNH Carnegie Museum of Natural History,

Invertebrate Zoology, 4400 Forbes
Avenue, Pittsburgh, Pennsylvania
15213 (Robert Davidson; davidsonr@
mus-nature.ca).

MAIC Michael A. Ivie Collection, Montana

State University, Department of
Biology, Bozeman, Montana 59717;
(Michael A. Ivie; mivie@montana.edu).
MCZC Harvard University, Museum of

Comparative Zoology, Department
of Entomology, 26 Oxford Street,
Cambridge, Massachusetts 02138—
2902 (Philip Perkins, perkins@oeb.
harvard.edu).

RHTC Robert H. Turnbow, Jr., Collection,

Directorate of Engineering and
Logistics, Fort Rucker, Alabama,
36362-5000 (Robert H. Turnbow,
Jr.; catch-bugs@aol.com).

USNM United States Department of

Agriculture, Systematic Entomology
Laboratory, c/o National Museum of
Natural History MRC 168, Wash¬
ington, D. C. 20560-0165 (Natalia J.
Vandenberg; nvandenb@sel.barc.usda.
gov).

WFBC William F. Barr Collection, 514

North Eisenhower Street, Moscow,
Idaho, 83844; Willian F. Barr; 208-
882-2886).

WOPC Weston Opitz Collection, Kansas

Wesleyan University, Department of
Biology, 100 East Claflin Avenue,
Salina, Kansas 67401-6196; Weston
Opitz; opitz@kwu.edu).

CHARACTERS SELECTED FOR
PHYLOGENETIC ANALYSIS

Fourteen anatomical characters serve to
establish an intrageneric phylogenetic hypoth¬
esis of Decorosa species and to root the genus
to its outgroup taxa. I relied on outgroup
comparisons to establish character polarity. In

AMERICAN MUSEUM NOVITATES

NO. 3628

this treatise, the outgroup involves members
of the genera Amboakis Opitz, Ellipotoma
Spinola, Hapsidopteris Opitz, Ichnea Laporte,
Katamyurus Opitz, Madoniella Pic, Megatra-
chys Opitz, Parvochaetus Opitz, and Plocam-
ocera Spinola. Character states identified as
“0” are judged plesiotypic whereas those given
a value of “1” are considered apotypic.
Derived character states marked by an asterisk
identify homoplasic apotypies, a type of
apotypy defined under “Materials and
Methods”.

Character 0 Seventh row of elytral punctations:

(0) present throughout elytral disc;
(1) absent in some portion of elytral
disc

Character 1 Elytral color pattern: (0) not multi¬
ples of streaks; (1) multiples of
streaks

Character 2 Labial terminal palpomere: (0) conic;
(1) curved-rectangular

Character 3 Pronotal anterior transverse depres¬
sion: (0) present; (1) absent*

Character 4 Pronotal contour: (0) convex; (1) flat

Character 5 Elytral punctations: (0) not binoded;
(1) binoded*

Character 6 Configuration of elytral posterior
margin; (0) obtuse; (1) acute*

Character 7 Eye size: (0) not small; (1) small

Character Vertex width: (0) not very wide; (1)
very wide*

Character 9 Phallic apex: (0) not hooked; (1)
hooked

Character 10 Form of sixth antennomere: (0) not
expanded; (1) expanded

Character 11 Number of antennomeres: (0) not 11

( 1 ) 10

Character 12 Phallic apex form: (0) not constricted
at middle; (1) constricted at middle

Character 13 Elytral disc markings: (0) not sharply
defined; (1) sharply defined

DECOROSA, NEW GENUS

Type Species: Decorosa iviei, n. sp.

Diagnosis: The known geographic distri¬
bution of the members of this genus is
confined to the Dominican Republic.
Morphologically, the most striking synapoty-
pic characteristic that distinguishes the mem¬
bers of this genus is that there has been a
reduction in expression of the seventh row of
elytral punctations. The seventh row is absent
in the anterior third or anterior half of the
elytral disc (see figs. 24, 29). Also, the
funicular antennomeres are filiform, and the
vertex is wider (fig. 2) than in specimens of
some of the superficially similar specimens of
Amboakis Opitz (2006: 22), Madoniella Pic
(1935: 10), and Parvochaetus Opitz (2006: 14).

Terminal palpomeres of the maxilla and
labium may also be used to distinguish
Decorosa specimens from those of other
above-mentioned epiphloeines. For example,
in Madoniella specimens the maxillary termi¬
nal palpomere is curved-subconic (fig. 36)
whereas in Decorosa specimens it is curved-
rectangulate (fig. 38). The labial terminal
palpomere is conic in specimens of
Madoniella (fig. 35), but slightly curved-conic
in those of Decorosa (fig. 37). The cranium
and pronotal disc are vested with short, pale,
horizontally decumbent, silken setae, the
elytral disc is very ornate with its pattern of
yellow and brown markings (fig. 1), and the
pronotum is longer than wide (fig. 7).

From superficially similar specimens of
the genera Madoniella, Amboakis, and
Parvochaetus, characterized by funicular anten¬
nomeres that are expanded (fig. 28), Decorosa
specimens are distinguished by less bulgy eyes
and a shallower medial border of the eye. Lastly,

TABLE 1

Character Matrix of 14 Morphological Characters

Taxa

Characters

outgroup -Decorosa stock

outgroup

aladecoris

limatula

iviei

neiba

2008

OPITZ: EPIPHLOEINAE, PART V: DECOROSA OPITZ

in specimens of Decorosa the epipleural fold
spans the entire length of the elytra (fig. 26),
which is not the case in specimens of Madoniella
(fig. 27), Amboakis, or Parvochaetus.

Description: Size: Length 3.5-5.6 mm;
width 1.0-1.4 mm. Form (fig. 12): Elongate,
narrow, rectangulate; pronotum elongate
(fig. 7) or about as long as wide (fig. 12);
elytra about three times longer than wide,
epipleural margin subparallel, posterolateral
margin gradually or sharply curved toward
sutural margin. Integument : Cranium and
pronotum light reddish brown to dark brown;
elytra with intermixture of yellow and brown
streaklike ornate patterns (figs. 1, 12-14); legs
yellow, tibiae may be infuscated at their
anterior margins; abdomen dark brown.
Vestiture : Cranium and pronotum vested
densely with short, pale, horizontally decum¬
bent, silken setae, elytral disc vested with
moderately stout, dark 1° setae, very uniform
in length, and fine decumbent pale 2°,
profusely distributed throughout elytral sur¬
face. Head : Cranium fine, rugosely punctate;
eyes not very bulgy, finely faceted, ocular
notch angle not acute at innermost point
(fig. 2); antenna (fig. 4) inserted at lower angle
of eye incision, comprised of 10 antennomeres,
loosely clubbed, scape about as long as
combined length of next three antennomeres,
pedicel oblong, funicular antennomeres fili¬
form, antennomeres 3-6 elongate, antenno-
mere7 quadrate, antennomeres 8 and 9 tria-
gonal, antennomerelO subglobose to ovate;
labrum (fig. 5) deeply incised, incision bor¬
dered by eight setae; mandible (fig. 10) robust,
anterior and posterior dentes well developed,
mandibular penicillus poorly developed; max¬
illa (fig. 9) well developed, terminal palpomere
curved-subconic, only slightly narrowed dis-
tally, laterolacinia present; labium (fig. 8) well
developed, terminal palpomere curved-rectan¬
gular; frons planar; gula (fig. 3) subtrapezoi-
dal. Thorax : Pronotum (figs. 6, 7) oblong or
about as long as wide, pronotal lateral
tubercle obtuse (fig. 12) or subacute (fig. 7);
anterior margin convex, posterior margin
linear, anterior transverse depression absent
(fig. 1) or short (figs. 12-14), disc slightly
depressed at sides where discal and paralateral
trichobothria are prominent; epimeral prolon¬
gations slightly extended to middle; procoxal

cavities open; elytra rectangulate, posterior
portion of epipleural margin gradually or
sharply rounded toward sutural margin, ante¬
rior half of disc surface macrosculptured with
nine rows of punctations, posterior half
macrosculptured with 10 rows of punctations,
seventh row of punctations absent in basal
half to basal third of disc (fig. 24); punctations
large, sometimes binodal (fig. 25), and much
wider than width of interstitial spaces; epi¬
pleural fold extends entire length of the elytra
(fig. 26), posterior rounded portion of epi¬
pleural margin minutely spinous; metathoracic
wing as in figure 23; mesoscutellum subqua¬
drate; tibial spur formula 0-1-1, tarsal pulvilli
formula 3-3-1, anterior margin of tibia with
one to three spines; tarsal claws with large
basal denticle. Abdomen: Six visible sterna;
pygidium broad scutiform. Male Genitalia:
Aedeagus short and broad, phallobasic apo-
deme short and broad, phallobasic rod usually
tapered to a fine point, phallic plates very
narrow; spicular fork as in figure 31.

Distribution: This insular genus is known
only from the Dominican Republic.

Etymology: The genus name stems from
the Latin adjective decorus (“beautiful”) and
the Latin feminine suffix -a. I refer to the
stunning elytral color pattern of these beetles.

KEY TO DIFFERENTIATE DECOROSA
FROM AMBOAKIS AND MADONIELLA,
AND KEY TO SPECIES OF DECOROSA

Epiphloeinae checkered beetles have four
trichobothria on the pronotum, two are
dorsal-paralateral (fig. 33) and one occurs
on each pronotal side. Also, the anterior
margin of the protibiae exhibit one to many
acute spines, which are not to be confused
with the more coarse and obtuse spines
found among some members of Enopliinae;
the latter tend to be in the 8-15 mm body-
size range whereas Decorosa specimens are
usually about 4 mm. This key deals only
with epiphloeines from Hispaniola.

1. Anterior half of elytral disc macrosculptured
by 10 rows of punctations, or elytral disc
somewhat roughened and densely matted
with decumbent setae to extent that
punctation organization poorly defined;

AMERICAN MUSEUM NOVITATES

NO. 3628

epipleural fold extended slightly beyond
middle of elytra (fig. 27); elytral disc vested

with bristlelike vertical setae.2

1'. Anterior half to anterior third of elytra
macrosculptured by nine rows of punctations
and by 10 behind middle, elytral disc never
roughened or densely matted with decum¬
bent setae; epipleural fold extended to elytral
apex (fig. 26); elytral disc not vested with

bristlelike vertical setae ( Decorosa ). 3

2(1). Funicular antennomeres subfiliform, fourth

and sixth slightly expanded (fig. 29); elytra
usually somewhat oblong-suboval (fig.

32). Madoniella Pic

2'. Funicular antennomeres very expanded

(fig. 28); elytra usually narrow-rectangu-

late (fig. 30). Amboakis Opitz

3(1'). Seventh row of elytral punctations begins at
about basal third of elytral disc, anterior to

the plane of metacoxae.4

3'. Seventh row of elytral punctations begins

near middle of elytral disc behind plane of

metacoxae (fig. 24). 5

4(3). Cranium and pronotum uniformly dark

brown; elytral markings as in figure 1

(Dominican Republic: La Vega).

. Decorosa aladecoris, n. sp.

4. Cranium and pronotum not uniformly dark

brown, cranium narrowly reddish, brown
at margins of eyes, pronotum narrowly red
brown along anterior margin (Dominican

Republic: San Juan).

. Decorosa neiba, n. sp.

5(3'). Pronotum narrow (figs. 13-14); apex of
anterior protibial margin of right tibia with
one stout spine; pronotal lateral tubercle
subacute (fig. 7); elytral disc markings as in
figures 13-14 (Dominican Republic: San

Juan: Santiago). Decorosa iviei, n. sp

5'. Pronotum broad (fig. 12); pronotal lateral
tubercle obtuse (fig. 12); apex of anterior
protibial margin of right tibia with 2
spines; elytral markings as in fig. 12

(Dominican Republic: Azua).

. Decorosa limatula, n. sp.

DESCRIPTION OF DECOROSA SPECIES

Decorosa aladecoris, new species
Figures 1, 34, 39, 42, 46

Holotype: Female. DOMINIC. REP.: La
Vega, Constanza, 1160 m, 30AUG1988, beat¬
ing in secondary pine guava forest, M. A. Ivie,
T. K. Philips & K. A. Johnson (USNM).
(Specimen mounted on on minuten pin in

neoprene block; gender label and metathorac-
ic wing mounted on support card; locality
label; holotype label).

Paratypes: Three specimens: Dominican
Republic: La Vega: Estacion Cabanito, 20-
VII-1996, R. Turnbow (RHTC, 1); Cordillera
Central, 4.1 km SW El Convento, 18°50'N
70°42'W, 31-IV-2003, 1729 m, disturbed ever¬
green forest with pine canopy trap, J. Rawlins,
R. Davidson, C. Young, C. Nunez, P. Acevedo
(CMNH, 1); Loma Casablito, 16.0 km NW
Bonao, 19°02°N 70°31'W, 28-IV-2003.1487 m,
evergreen cloud forest at summit, canopy trap,
J. Rawlins, R. Davidson, C. Young, C. Nunez,
P. Acevedo (WOPC, 1).

Diagnosis: From congeners these beetles
are distinguished by the uniform dark brown
coloration of the cranium and pronotum, and
by the color pattern on the elytral disc (fig, 1).

Description: Size : Length 4.0 mm; width
1.2 mm. Integumental Color. Cranium and
pronotum dark brown; antennae yellow;
elytra as in figure 1; legs yellow; venter dark
brown. Vestiture: Cranium and pronotum
densely vested with pale decumbent setae.
Head : Vertex much wider than width of eye
(27:13); ratio of head width to pronotal width
1.05 (62:59); antennal form similar to antenna
depicted in figure 4. Thorax : Pronotum only
slightly longer than broad (60:58); pronotal
lateral tubercle obtuse; anterior margin con¬
spicuously wider than posterior margin
(58:50); apical region of protibial anterior
margin with 3 spines; seventh row of elytral
punctations begins slightly anterior to elytral
middle. Abdomen : Aedeagus as in figure 34.

Variation: There is some variation in the
extent of dark and light coloration in the most
distal region of the elytral disc.

Natural History: Specimens were col¬
lected during May, July, and August, one by
beating secondary vegetation in a pine/guava
forest at 1160 m. Two specimens were cap¬
tured in canopy traps, one in disturbed
evergreen forest with pine at 1729 m, the
other at the summit of an evergreen cloud
forest at 1487 m.

Distribution (fig. 46): Known only from
the highlands of the La Vega Province of the
Dominican Republic.

Etymology: The specific epithet is a Latin
compound name that stems from the noun ala

2008

OPITZ: EPIPHLOEINAE, PART V: DECOROSA OPITZ

Fig. 1. Habitus of Decorosa aladecoris Opitz.

AMERICAN MUSEUM NOVITATES

NO. 3628

(“wing”) and decoris (“adorned”). I refer to
the ornate markings on the elytral disc.

Decorosa iviei, new species
Figures 2-11, 13, 14, 19-26, 37-38, 46

Holotype: Female. DOMIN. REP: Prov.
San Juan, Pico Duarte weather sta., 2885 m,
07APR.-08JUL 1992, lindgren funnel, M. A.
& R. O. I vie (USNM). (Specimen point
mounted, gender label affixed to paper point;
support card; locality label; holotype label.)

Paratypes: Seven specimens. Dominican
Republic: La Vega: Loma Rucilla & moun¬
tains, N, day not noted-VI-1938, 1524-
2439 m, Darlington (MCZC, 2; AMNH, 1);
53 km SE Constanza, 9-VIII-1979, L. B.
O’Brien (WFBC, 1); Parque Nacional
Armando Bermudez, N Pico del Yaque, 29-
VII-1993, 2375 m, beating vegetation, D.
Sikes & R. P. Rosenfeld (MAIC, 2; WOPC, 1).

Diagnosis: The more narrow pronotum,
more angular lateral pronotal tubercle, singu¬
lar spine on the anterior margin of the
protibia, and configuration of the elytral
pattern (figs. 13, 14) conveniently distinguish
the members of this species from the superfi¬
cially similar specimens of Decorosa limatula.

Description: Size: Length 3.8-5.8 mm;
width 1.0-1.3 mm. Integumental Color: Cra¬
nium and pronotum light reddish brown;
antennae yellow; elytra as in figure 13 and
14; legs predominantly yellow, tibiae infus-
cated; venter dark brown. Vestiture: Cranium
and pronotum densely vested with pale
decumbent setae. Head: Vertex about four
times wider than width of eye (12:40); ratio of
head width to width of pronotum 1.18 (67:57);
antennae as in figure 4. Thorax: Pronotum
only slightly longer than broad (60:58);
pronotal lateral tubercle acute; side margins
very sinuous; anterior margin conspicuously
wider than posterior margin (56:47); apical
region of protibial anterior margin with one
spine. Abdomen: Aedeagus as in figures 19-22.

Variation: In one specimen the cranium
and pronotum is dark brown rather than
reddish brown and the expression of light and
dark coloration of the elytral disc varies as
indicated in figures 13 and 14. These figures
represent the extremes in elytral color density
among the species examined.

Natural History: Members of this spe¬
cies have been captured during June-August
at altitudes ranging from 1524-2885 m.

Distribution (fig. 46): This species is
known from the highlands of the Cordillera
Central of the Dominican Republic.

Etymology: The trivial name is a dedica-
tive patronym to express appreciation to
Michael A. Ivie for his contributions to West
Indian coleopterology.

Decorosa limatula, new species
Figures 12, 15-18, 46

Holotype: Female. Loma Vieja, S Con-
stanza, Aug. ’38, Dom. Rep., c. 6,000 ft.,
Dari. (MCZC). (Specimen point mounted,
gender label affixed to paper point; support
card; locality label; holotype label.)

Paratypes: None.

Diagnosis: The broadly flattened prono¬
tum, less acute lateral pronotal tubercle,
bispinous condition of the anteroapical mar¬
gin of the protibia, and configuration of the
elytral pattern (fig. 12) readily distinguish the
members of this species from congeners.

Description: Size: Length 5.0 mm; width
1.3 mm. Integumental Color: Cranium and
pronotum light red-brown, vertex infuscated;
antennae yellow; elytra as in figure 12; legs
predominantly yellow, tibiae infuscated; ven¬
ter dark brown. Vestiture: Cranium and
pronotum densely vested with pale decumbent
setae. Head: Vertex about two times wider
than width of eye (19:34); antennae generally
as in figure 4. Thorax: Pronotum only slightly
longer than broad (70:68), broadly flattened;
pronotal lateral tubercle obtuse; side margins
not very sinuous; anterior margin conspicu¬
ously wider than posterior margin (65:60);
protibial anteroapical region with two spines.
Abdomen: No aedeagal information available.

Variation: One specimen examined.

Natural History: The unique specimen
was collected in August at 1829 m.

Distribution (fig. 46): This species is
known from the highlands of south-central
Dominican Republic.

Etymology: The specific epithet is a Latin
adjective derived from limatulus (“rather
polished”) with a feminine ending to modify
the genus name. I refer to the shimmering

Figs. 2-11. Anatomical organs of Decorosa iviei Opitz. 2. Head (anterior view). 3. Head (posterior view).
4. Antenna. 5. Labrum. 6. Pronotum (ventral view). 7. Pronotum (dorsal view). 8. Labium. 9. Maxilla. 10.
Mandible. 11. Metendosternite. Scale bars in all figures represent 1 millimeter.

AMERICAN MUSEUM NOVITATES

NO. 3628

Figs. 12-22. Habiti and aedeagi. 12. Habitus, Decorosa limatula Opitz. 13-14. Habiti, D. iviei Opitz. 15-
18. Aedeagus, D. limatula (15, tegmen, ventral view; 16, tegmen, lateral view; 17, phallus, dorsal view; 18,
phallus, lateral view). 19-22. Aedeagus, D. iviei (19, tegmen, ventral view; 20, tegmen, lateral view; 21,
phallus, dorsal view; 22, phallus, lateral view).

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OPITZ: EPIPHLOEINAE, PART V: DECOROSA OPITZ

Figs. 23-27. Anatomical organs. 23-26. Decorosa iviei Opitz (23, metathoracic wing; 24, elytra, ventral
view; 25, cluster of elytral punctations; 26, elytra, lateral view). 27. Madoniella dislocata (Say) elytra, lateral view.

characteristic that the short pale decumbent
setae bestow on the cranium and pronotal disc.

Decorosa neiba, new species
Figures 40, 41, 43, 46.

Holotype: Female. DOMINICAN RE¬
PUBLIC: Elias Pina. Cierra de Neiba,
9.1 km WSW Hondo Valle, 18-41-38N, 71-
46-56W, 1856 m, 25 June 2003, J. Rawlins, C.
Young, R. Davidson, C. Nunez, P. Acevedo,

M. de la Cruz, wet montane forest with pine,
malaise trap, Carnegie Museum Specimen
Number CMNH-356.509 (CMNH). (Specimen
minuten pin mounted on neoprene block;
gender label and metathoracic wing mounted
on support card; locality label; holotype label).

Paratypes: One specimen: Dominican
Republic: San Juan: Sierra de Neiba, 9.4 km
SSW El Cercado, 18° 39'N 71° 32'W, 22-VI-
2003, 1974 m, mature pine forest, Malaise
trap, R. Davidson, C. Nunez, C. Young, J.
Rawlins, P. Acevedo, M. de la Cruz (WOPC, 1).

AMERICAN MUSEUM NOVITATES

NO. 3628

Figs. 28-32. Anatomical structures and body outlines. 28. Amboakis nova (Opitz) antenna. 29.
Madoniella dislocata (Say) antenna. 30. A. nova body outline. 31. Decorosa iviei Opitz spicular fork. 32. M.
dislocata body outline.

Diagnosis: From superficially similar
specimens of D. aladecoris, D. neiba specimens
may be distinguished by the shape of the
pronotum. Specimens of A. neiba have a
proportionally wider pronotal posterior mar¬
gin (compare figs. 42, 43). From other conge¬
ners these beetles are distinguished by the
bicolorous coloration of the pronotum; the
disc is dark brown and the anterior margin is
reddish brown.

Description: Size : Length 4.0-4.2 mm;
width 1.6-1.8 mm. Integumental Color.
Cranium and pronotum bicolorous, cranial
disk dark brown, periphery reddish brown,
pronotal disc dark brown, anterior margin
reddish brown; antennae unicolorous, mostly
yellow, some funicular articles infuscated;
elytra as in figure 41; legs mostly yellow,

femora and tibiae infuscated; venter dark
brown. Vestiture: Cranium and pronotum
densely vested with pale decumbent setae.
Head : Vertex much wider than width of eye
(13:25); ratio of head width to pronotal width
1.06 (57:54); antennal form similar to antenna
depicted in figure 4. Thorax : Pronotum only
slightly longer than broad (53:52); pronotal
lateral tubercle obtuse; anterior margin slight¬
ly wider than posterior margin (48:47); apical
region of protibial anterior margin with three
spines; seventh row of elytral punctations
begins slightly anterior to elytral middle.
Abdomen : Aedeagus as in figure 40.

Variation: The available specimens are
quite homogeneous.

Natural History: The two available
specimens were collected in a Malaise trap in

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OPITZ: EPIPHLOEINAE, PART V: DECOROSA OPITZ

Figs. 33-38. Anatomical structures. 33. forebody of Madoniella dislocata (Say). 34. Aedeagus of
Decorosa aladecoris Opitz. 35-36. Madoniella dislocata (35, terminal labial palpomere; 36, terminal maxillary
palpomere). 37-38. D. iviei Opitz (37, terminal maxillary palpomere; 38, terminal labial palpomere.

AMERICAN MUSEUM NOVITATES

NO. 3628

Figs. 39^13. Anatomical organs and habitus. 39. Decorosa aladecoris Opitz elytra, ventral view. 40-41.
D. neiba Opitz (40, aedeagus; 41, habitus). 42. D. aladecoris , pronotum. 43. D. neiba, pronotum.

mountain forest laden with pine, at 1857 and
1974 m.

Distribution (fig. 46): Known only from
the highlands of Sierra de Neiba, in the San
Juan Province of the Dominican Republic.

Etymology: The trivial name, neiba, con¬
stitutes a noun in apposition and refers to the
type locality.

REVIEW OF GREATER ANTILLEAN AND
HISPANIOLOAN PALEOGEOGRAPHY

The literature is rich in facts, ideas, and
syntheses about Caribbean biogeography/

geology (Schwartz, 1980; Rosen, 1985;
Liebherr, 1988; Mann et al., 1991; Donovan
and Jackson, 1994; Woods and Sergile, 2001,
and references therein) and more specifically
about insect Hispaniolan zoogeography such
as Woodruff and Sanderson (2004: 26). These
stimulating contributions inspire attempts to
make biographical sense of one’s taxonomic
findings. My intent for this review of Antillean
paleohistory is to provide a baseline of
thoughts, along with additional comments
related to insect findings, and summary for
this and future analyses of Caribbean Cleridae
taxa.

2008

OPITZ: EPIPHLOEINAE, PART V: DECOROSA OPITZ

.60

■8

■S

-10

-13

-■12

-9

/—outgroup-Decorosa stock

.<d

-6

-7

■3

-4

-8

-0

-1

-2

outgrou p-Decorosa stock

Fig. 44. Phylogenetic tree of Decorosa Opitz
and related taxa; manually prepared.

There is strong evidence from geology
(Khudoley and Meyerhoff, 1971: 129; van
Fossen and Channell, 1988: 610; Perfit and
Williams, 1989: 67; Heubeck et.al., 1991: 29;
Draper et al., 1994: 143) and biology

(Williams, 1961: 2; Schwartz, 1980: 87, 1989:
489; Rosen, 1985: 652; Liebherr, 1988: 143;
Woodruff and Sanderson, 2004: 27) to sup¬
port the hypothesis that Hispaniola was once
comprised of two islands, and that the
northern part originated from an early
Cretaceous-Eocene assemblage of insular rock
whereas the southern portion originated from
oceanic plateau rock, essentially the uplifted
edge of Caribbean sea crust (Draper et al.,
1994: 129). The two-island hypothesis and the
prehistoric movement of the two landmasses
that constitute Hispaniola have been incorpo¬
rated into a paleogeographic model of West
Indies mobility (Pindell and Dewey, 1982: 202;

-I — outgroup

h o r o /w w v—

wt ( 9 9 1

1111

6 7 9

— limatula

Fig. 45. Phylogenetic tree of Decorosa Opitz
and related taxa; computer generated.

Sykes et al., 1982: 10,669; Wage and Burke,
1983: 633; Rosen, 1985: 652; Perfit and
Williams, 1989: 67; Pindell and Barrett, 1990:
405).

To explain biologic relationships between
the northern and southern elements of
Hispaniola, and those between Hispaniola
and any other West Indies landmass, we must
explore what is known about the geology and
the paleotectonic and neotectonic events that
eventually led to the island’s amalgamation.
Moreover, the relative geologic ages of the two
prehistoric landmasses is relevant to any
discussion of their biogeography.

Donnelly (1988: 26) places the initial emer¬
gence of Greater Antillean terrain during the
upper lower Cretaceous, about 105 Ma with
Hispaniolan and Jamaican rocks included in
the mix. Khudoley and Meyerhoff (1971: 152)
suggest that “Most of the Greater Antilles
were uplifted in the Miocene and later times.”
Early Cretaceous rock elements in northern
Hispaniola and late Cretaceous rock elements
for the southwestern part of the island were
reported by Draper et al. (1994: 129). This
suggests that at least some portions of the
crustal component of the Hispaniolan north¬
ern island may be more ancient than its
southern counterpart. Rocks of the late upper
Cretaceous (Maastrichtian stage, about
70 Ma) have been reported as a component
of the southwestern Sierra de Bahoruco
(Draper et al., 1994: 139). Of considerable
relevance to the question of age of the
southern portion of Hispaniola is the gymno-
sperm and angiosperm fossil discovery by
Woodruff (1986: 32; also see Woodruff and
Fritsch, 1989: 216) from the Larimer deposits

AMERICAN MUSEUM NOVITATES

NO. 3628

of the Sierra de Baoruco considered Creta¬
ceous by De Leon (1989: 85).

It has been hypothesized that Hispaniola’s
southern terrain was in proximity with the
Yucatan peninsula (Rosen, 1985: 644) during
early Eocene to late Paleocene (from 50-
65 Ma) and that crustal movements, involving
fault generated tectonics, between the
Caribbean and North American plates carried
southern Hispaniola to its eventual welding
with the northern Hispaniolan component; an
amalgamation purported to have occurred
during early Miocene (about 20 Ma)
(Heubeck et al., 1991: 29). Depending on
one’s starting point, this means that the two
islands have been separated from 35 to
85 million years of Caribbean evolution.
And, while both islands experienced tumultu¬
ous geologic changes (Donnelly, 1988: 27),
likely environmental stresses from a bolide
impact (Hedges, 2001: 19; Opitz, 2005: 108),
and presumably vicissitudinous climates
(Clench, 1963: 257), which must have had
significant impacts on speciation and extinc¬
tions (Curtis et al., 2001: 50; Morgan, 2001:
398), the aforementioned time span represents

a wide window of opportunity for independent
biologic evolution.

Moreover, during their eastward tectonic
journey, the terrestrial components of
Hispaniola would have experienced the kind
of fault-based shifting and rotation that would
have placed them in close geographic relation¬
ships with other Greater Antillean terrain.
Draper and Barros (1988: 60, 1994: 82)
suggest late Eocene geologic ties between
Cuba and Hispaniola and based on
Lygaeidae findings Baranowski and Slater
(1998: 75) suggests vicariance between Cuba
and Hispaniola, and possibly Puerto Rico. In
a treatise of plant phylogeny involving trees
and shrubs of Lyonia (Ericaceae), Judd (2001:
72) discusses geologic relationships between
western Cuba and southwestern Hispaniola
and biotic relationships among Puerto Rico,
north-central Hispaniola, and eastern Cuba.

If the welding of northern and southern
Hispaniola occurred some 20 Ma (Heubeck et
al., 1991: 29) and speciation events requires
about 3 million years, as suggested by
Whitehead (1972: 308) in his work with
ground beetles, then biologists interested in

2008

OPITZ: EPIPHLOEINAE, PART V: DECOROSA OPITZ

Hispaniolan beetle evolution have some mea¬
sure for the temporal character of lower taxa
divergence. This would be particularly useful
for temporal analyses involving sister-group
relationships between northern and southern
Hispaniola and vicariant relationships among
taxa from Hispaniolan and those of other
Greater Antillean terrain.

DECOROSA EVOLUTIONARY
CONSIDERATIONS

Two phylogenetic trees (figs. 44 and 45)
were produced for Decorosa, one (fig. 44)
prepared “by hand” and the other (fig. 45)
computer generated. The computer analysis
generated one parsimonious tree of 29 steps,
Cl of 100, and an RI of 100. Both phylogenies
produced identical pairs of Decorosa sister
species.

Paucity of Decorosa specimens and few
distributional records usually preclude com¬
prehensive discussion about phylogeny and
zoogeography. But even a few tentative
comments might be beneficial for future
endeavors about the subject. Hispaniola has
benefited from a flurry of entomological
activities in recent years (Schwartz, 1989:
489; Woodruff and Sanderson, 2004: 1; and
various communications with insect collec¬
tors) and there has evolved a greater aware¬
ness of insect diversity in the Caribbean
(Liebherr, 1988). Some of these activities
include greater insect collecting efforts, which
have resulted in greater availability of Cleridae
specimens.

The four Decorosa species are known only
from the northern highlands of Hispaniola
(fig. 45) with D. limatula, the sister species of
the more northern D. iviei found in highlands
east of Sierra de Neiba slightly north of the
Enriquillo basin. Decorosa aladecoris and D.
neiba, the more primitive members of the
genus, are undoubtedly sympatric with the
most derived D. iviei, which at the very least
suggests some dispersal element in the early
history of the genus.

As D. limatula is known from environs
slightly north of the Enriquillo basin, one
might presume a vicariant relationship be¬
tween D. limatula with the more northern
sister species D. iviei, as is the case between D.

aladecoris and D. neiba. Discovery of limatula
specimens from south of the Enriquillo basin
would certainly support that possibility, and
corroborate the likely north-south vicariance
findings of Mertens (1939: 12), Williams
(1961: 2), Schwartz (1980: 90), Judd (2001:
69), and Woodruff and Sanderson (2004: 26).
As the most primitive members of the genus
were collected from the northern portion of
Hispaniola, I suspect that the northern com¬
ponent of the two paleoislands served as the
ancestral terrain of the genus in which
ancestral Decorosa stock B diverged to pro¬
duce the aladecoris-neiba lineage and the more
derived limatula-iviei lineage (fig. 44). More
widespread northern collections of additional
D. aladecoris and neiba specimens or absence
of such material from the southern island
would support the hypothesis and more
credibly root the initial geographic divergence
of the genus. Sister-group relationships of
insect taxa between these two formerly
insular highlands have been confirmed in
Scarabaeidae (Woodruff and Sanderson,
2004: 26) and in butterflies (Schwartz, 1989:
489; Sourakov, 2000: 79).

The putative absence of Decorosa from
western and southwestern Hispaniola could
be a consequence of collecting bias or simply
that captured specimens are unknown to me.
I, like many other biologists, fear that
extensive deforestation has eliminated a sub¬
stantial Hispaniolan insect fauna, especially
from western portions of the island (Sergile
and Woods, 2001: 547).

The divergence of ancestral Decorosa yield¬
ed the aladecoris line, which retained several
primitive characteristics including normal eye
size and nonnodal elytral punctations while
acquiring an apotypic convex pronotum and
loss of the pronotal anterior transverse de¬
pression. The sister taxon of this lineage,
ancestor C, would have evolved a more
flattened pronotum, the eyes became signifi¬
cantly reduced, and the elytral punctations
acquired a binodal characteristic. This ances¬
tor eventually yielded D. limatula and D. iviei
each characterized as indicated in fig. 44. It
remains to be seen whether D. limatula
dispersed to southern portions of prehistoric
Hispaniola from northern ancestral grounds,
or whether the north-south distribution record

AMERICAN MUSEUM NOVITATES

NO. 3628

of the limatula-iviei lineage expresses some
manner of disjunction between the paleois-
lands of Hispaniola.

ACKNOWLEDGMENTS

I am indebted to Lee Herman Jonathan R.
Mawdsley, and Robert E. Woodruff for their
review and improvement of the manuscript.
As always, Jean-Michel Maes generously
translated the abstract into Spanish. My
thanks to Greg Zolnerowich for the photo¬
graph of D. aladecoris and to the keepers of
collections for trust, patience, and other loan
courtesies. My sincere appreciation to Mary
Knight (AMNH) for her editorial work, which
substantially improved the manuscript.

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Complete lists of all issues of the Novitates and the Bulletin are available at World Wide
Web site http://library.amnh.org/pubs. Inquire about ordering printed copies via e-mail from
scipubs@amnh.org or via standard mail from: American Museum of Natural History,
Library—Scientific Publications, Central Park West at 79th St., New York, NY 10024. TEL:
(212) 769-5545. FAX: (212) 769-5009.

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Full text of "Classification, natural history and evolution of the Epiphloeinae (Coleoptera, Cleridae). Part 5, Decorosa Opitz, a new genus of checkered beetles from Hispaniola with description of its four new species"

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