ALABAGRUS ENDERLEIN (HYMENOPTERA, BRACONIDAE, AGATHIDINAE) SPECIES OF COSTA RICA, WITH AN EMPHASIS ON SPECIMENS REARED FROM CATERPILLARS IN AREA DE CONSERVACIÓN GUANACASTE

2018·32 MB·

by  Michale J Sharkey

18 June 2018
Contributions in Science, 526:31(cid:8212)180
ALABAGRUS ENDERLEIN (HYMENOPTERA, BRACONIDAE, AGATHIDINAE)
SPECIES OF COSTA RIcA, WITH AN EMPHASIS ON SPECIMENS REARED
FROM CATERPILLARS IN AREA DE CONSERVACION GUANACASTE(cid:8221)
BOHibela): SHARKEY,(cid:8221) > SARAH MEIEROTTO,(cid:8221) ERIC CHAPMAN,(cid:8221) DANIEL
JANZEN,(cid:8221) WINNIE HaLLWACHS,(cid:8221) TANYA DAPKEY,(cid:8221) AND M. ALMA Sous*
ABSTRACT. Eighty-six species of Costa Rican Alabagrus are treated; these include species reared from lepidopteran larvae in Area de
Conservacion Guanacaste, Costa Rica, over 32 years of caterpillar inventory. Sixty-five new species are described, that is, A. almasolisae, A.
andresfreitasi, A. andywarreni, A. axelhausmanni, A. barbsharanowskiae, A. bernardoespinozai, A. bobpoolei, A. bobrobbinsi, A. bobwhartoni, A.
brendameierottoae, A. brianbrowni, A. brianharrisi, A. craigevansi, A. donharveyi, A. donlafontainei, A. fernandezi, A. fernandodiasi, A.
genemonroei, A. hansoni, A. hespenheidei, A. iankitchingi, A. ilgookangi, A. isidrochaconi, A. jackiemillerae, A. jeanfrancoislandryi, A
jeanmariecadioui, A. jennyphillipsae, A. jimmilleri, A. johnbrowni, A. johnburnsi, A. johnobryckii, A. johnsharkeyi, A. kaciejoae, A. karensharkeyae,
A. kaydodgeae, A. keithwillmotti, A. leedyeri, A. lindapitkinae, A. longinoi, A. malcolmscoblei, A. marcepsteini, A. mariaheikkilae, A. markmetzi, A
mattottoi, A. nickgrishini, A. patsharkeyi, A. paulgoldsteini, A. paulheberti, A. paulsharkeyi, A. paulthiaucourti, A. quickei, A. ramyamanjunathae, A
reddypallii, A. rudolfmeieri, A. sarahmeierottoae, A. sarahsharkeyae, A. scottmilleri, A. scottshawi, A. semihespenheidei, A. stiremani, A.
tanyadapkeyae, A. tommyersi, A. victoriapookae, A. yuanmaofangi, and A. yuchinkengae.
Species limits primarily based on cytochrome c oxidase subunit I mitochondrial DNA sequence data greatly differed from previous
morphological attempts to delimit species of Alabagrus. As a result, 17 species reported to occur in Costa Rica are no longer considered to be
present in the country, that is, A. albispina, A. imitatus, A. juchuy, A. kagaba, A. latisoma, A. latreillei, A.m aya, A. mojos, A. nahuatl, A. nigrilitus,
A. pachamama, A. paruyana, A. parvifaciatus, A. semialbus, A. tricarainatus, A. tripartitus, and A. warrau. Furthermore, five species have been
found to be composed of species complexes, that is, A. cocto, A. englishi, A. pecki, A. roibasi, and A. yaruro. These difficulties point to the
impossible task of delimiting species of Agathidinae solely with morphological evidence.
An illustrated key to species, and a plate of color photos of each species are provided.
INTRODUCTION as the outgroup for the phylogenetic analyses. Both genera are
restricted to the New World.
This article is the third in a series revising the species of Agathidinae We include a molecular data set consisting of 680 Alabagrus
reared from lepidopteran caterpillars as part of the inventory of the operational taxonomic units (OTUs) sequenced for cytochrome c
caterpillars and their parasitoids of Area de Conservacién Guanacaste oxidase subunit I (CO/) for approximately 60 species. The majority of
(ACG), northwestern Costa Rica (http://janzen.sas.upenn.edu; http:// specimens (624) were reared from caterpillars (those with specimen
www.acguanacaste.ac.cr; Janzen et al., 2009). It includes a key to 87 numbers beginning with DHJPAR).
species of Alabagrus Enderlein, of which 65 are newly described. This Alabagrus was first revised by Sharkey (1988), who included 104
is not meant to be a comprehensive treatment of the Costa Rican
species. Leathers and Sharkey (2003) revised the species of Alabagrus of
fauna of Alabagrus. A conservative estimate, based on extensive La Selva, and included previously described species known to occur in
collections at Instituto Nacional de Biodiversidad (INBio; Costa Costa Rica. They treated 39 putative species, including six new species.
Rica), the Universidad de Costa Rica, and the Hymenoptera Institute These two revisions relied solely on morphological evidence. The species
(Lexington, Kentucky), is that about 300 species occur in the concepts in the present treatment were formulated with the assistance of
country. COI sequence data and a plethora of host data as the result of rearings in
New World members of Al/abagrus may be distinguished from other Area de Conservacién de Guanacaste. These data have had a profound
agathidines by the following combination of character states: simple effect. There are many more cryptic species than previously thought,
claws with a basal lobe, frons bordered laterally by carinae; first and many species with presumed wide distributions are more likely to be
metasomal median tergite distinctly convex and/or with a median species complexes whose component species have relatively restricted
ridge, gena not elongate. All are presumed to be koinobiont distributions. As a result, 17 species, reported to occur in Costa Rica by
endoparasitoids of concealed larvae of Crambidae. Pharpa Sharkey, Leathers and Sharkey (2003), are no longer considered to be present in
1988, is the sister group to Alabagrus (Sharkey, 1988) and is used here the country, these are Alabagrus albispina, A. imitatus, A. juchuy, A
kagaba, A. latisoma, A. latreillei, A. maya, A. mojos, A. nahuatl, A.
(cid:8216)URL: www.nhm.org/scholarlypublications
nigrilitus, A. pachamama, A. paruyana, A. parvifaciatus, A. semialbus, A
* Department of Entomology, University of Kentucky, $-225 Agricultural
Science Center North, Lexington, Kentucky 40546-0091, USA. tricarainatus, A. tripartitus, and A. warrau. Furthermore, five of the
z Department of Biology, University of Pennsylvania, 3740 Hamilton species reported by Leathers and Sharkey (2003) have been found to be
Walk, Philadelphia, Pennsylvania 19104, USA.
composed of species complexes, that is, Alabagrus cocto, A. englishi, A
Systematic Entomology Laboratory, Agriculture Research Service, U.S.
pecki, A. roibasi, and A. yaruro. These difficulties point to the impossible
Department of Agriculture, Smithsonian Institution National Museum of
task of delimiting species of Agathidinae solely with morphological
Natural History, E-517, MRC 168, Washington, D.C. 20013-7012, USA.
> Corresponding author: Michael J. Sharkey, E-mail: [email protected] evidence.
© Natural History Museum of Los Angeles County, 2018
ISSN 0459-8113 (Print); 2165-1868 (Online)
32 HE Contributions in Science, Number 526 Sharkey et al.: Alabagrus of Costa Rica
METHODS MORPHOLOGICAL TERMS
Morphological terms are from Sharkey and Wharton (1997). The reader
SPECIMEN INFORMATION
is advised to search for any morphological term that is not understood
Most specimens used in this study are deposited in the Entomology
on the Hymenoptera Anatomy Ontology website (http://portal.hymao.
Museum of Utah State University and the Hymenoptera Institute
org/projects/32/public/ontology/; Yoder et al., 2010). To do this, enter
Collection, with the exception of specimens with numbers beginning the term in the search box, select the best match from the list that pops
with (cid:8220)BIOUG,(cid:8221) which are deposited at the University of Guelph,
up, and click on the show button.
Centre for Biodiversity Genomics, Ontario, Canada. Source files for the
keys, descriptions, illustrations, DNA sequences and distributional data MUSEUM ACRONYMS
are all freely available to future researchers who may wish to build on
these records. The detailed specimen records are available by search of CNCI Canadian National Collection of Insects, Ottawa, Canada
EMUS Utah State University, Department of Biology, Entomology
the individual specimen DHJPARxxxxxxx voucher codes at the Janzen
database (http://janzen.bio.upenn.edu/caterpillars/database.lasso). Some Museum, Logan, Utah, USA
HIC University of Kentucky, Department of Entomology, Hymenop-
of the Lepidoptera hosts are incompletely identified; however, they also
tera Institute Collection, Lexington, Kentucky, USA
have unique names, such as Desmia Solis19 (which is an interim name
INBio Instituto Nacional de Biodiversidad, Santo Domingo de Heredia,
for Desmia species 19 as determined by M. Alma Solis of the U.S.
Costa Rica
Department of Agriculture Systematic Entomology Laboratory, Wash-
MUCR Ciudad Universitaria, Universidad de Costa Rica, Museo de
ington, D.C.). These names will be updated in the Janzen database
when the species is baptized with a formal scientific name, but the Insectos, Costa Rica
interim name, in this case Desmia Solis19, will remain searchable in that
database. Host caterpillars are uniquely identified by their own voucher
DNA EXTRACTION, PCR, AND SEQUENCING
code system, which is recognizable by YY-SRNP-XXXXX, where YY is
Specimens sequenced in this study are from a variety of sources.
the two-digit year and XXXXX is a unique number within that year.
Specimens with numbers beginning with DHJPAR, INB, INBIOCRI,
DNA trace files and primer information are available through the
or BIOUG were extracted and sequenced for COJ at the University of
Barcode of Life Data Systems (BOLD, http://www.boldsystems.org;
Guelph following the methods outlined in Hebert et al. (2004).
Ratnasingham and Hebert, 2007), as is specimen information for
Specimens with numbers beginning with the letter (cid:8220)H(cid:8221) were
specimen numbers INBxxxxxxxxxx, INBIOCRIxxxxxxxxx, and BIO-
extracted from individual legs with the QIAGEN DNeasy Blood and
UGxxxxx-xxx. Specimen information for specimens with numbers
Tissue Kit using the animal tissue protocol (QIAGEN Inc., Chatsworth,
beginning with the letter (cid:8220)H(cid:8221) are stored in the Symbiota database
California, USA). The mitochondrial COJ (~658 bp) gene was
(Gries et al., 2014; Symbiota Collections of Arthropods Network,
amplified with the primer pair LepFl and LepR1 (Hebert et al.,
http://symbiota4.acis.ufl.edu/scan/portal/) under the Hymenoptera
2004). Polymerase chain reaction (PCR) was conducted using TaKaRa
Institute Collection (HIC). To search for a specimen in the database,
reagents, with each reaction consisting of 1X buffer, 0.3 mM
do the following: Under the Search tab (upper left of the screen), select
nucleotides, 0.4 tM of each primer, 0.625 U TaKaRa Ex Taq,
Search Collections. Deselect all collections, and scroll down the Southeast
double-distilled water, and 1-3 tL template DNA in a total reaction
section, put a check in the box next to Hymenoptera Institute Collection,
volume of 25 wL. The thermal cycling protocol had an_ initial
scroll back up and hit the Search button (right side of screen). H-
denaturation period at 95 °C for 2.5 min, followed by 40 cycling steps
numbered specimens are stored with a four-letter prefix (HICH)
which denatured at 95 °C for 30 s, annealed at 44 °C for 30 s and
followed by a six-digit number. Therefore, H369, as it appears in this
extended at 68 °C for 45 s, with a final extension step of 72 °C for 7
publication, is stored as HICH000369. To search for this specimen,
min. To determine reaction success, PCR products were electrophoresed
scroll down to the Specimen Criteria section, type this number in the
in 1% agarose stained with ethidium bromide. PCR products were
box next to Catalog Number and hit the search button. This displays a
outsourced for Sanger sequencing either by the Advanced Genetic
page with a summary of the specimen information. Clicking on Full
Technologies Center (University of Kentucky, Lexington, Kentucky,
Record Details opens a new window with the full specimen record,
USA) or Beckman Coulter Genomics (Danvers, Massachusetts, USA)
including all available images.
using labeled dideoxy-nucleotides with ABI 3730, Big-Dye Terminator
Each species is diagnosed and described or redescribed, and a plate of
mix v. 3.0, or with ABI PRISM 3730xl, BigDye Terminator mix v. 3.1
images with a chronogram of occurrence is included for each.
(Applied Biosystems, Foster City, California, USA).
SPECIES CONCEPTS
DNA ASSEMBLY AND PHYLOGENETIC ANALYSIS
The biological species concept is used with evidence drawn from COJ
Bidirectional sequences were aligned and edited using Geneious Pro (v.
sequences, morphology, and host use as indirect evidence for species
6.1.5; Drummond et al., 2009) and multiple alignments were assembled
integrity. COJ was the primary source of evidence with no particular
using MAFFT (v. 5; Katoh et al., 2006) using the default settings and
percent divergence employed as a cut-off. The sequence data were
refined by eye using MacClade v. 4.08 (Maddison and Maddison,
checked against morphological and host-use data, and these corrobo- 2005). Maximum likelihood (ML) phylogenetic analyses were conduct-
rated or (rarely) refuted the sequence data.
ed on the CO/ data set using Garli (v. 2.01; Zwickl, 2006). The data set
consisted of 680 Alabagrus OTUs and one sequence from a species of
TAXON SAMPLING
Pharpa Sharkey, which was shown to be sister to A/abagrus in the most
For molecular analyses, we used all Alabagrus sequences available to us comprehensive molecular agathidine phylogeny to date (Sharkey and
on the BOLD database from Costa Rica. Once we removed a few Chapman, 2017) for outgroup rooting. The data were partitioned by
sequences that were contaminated in some way, this amounted to 636 codon position (three partitions). We applied the most complex model
sequences. To this, we added 44 Alabagrus sequences generated by the available (GTRHHG; Rodriguez et al., 1990) to each partition as per
Sharkey lab from Costa Rica, Honduras, Mexico, and USA. recommendations of Huelsenbeck and Rannala (2004) for likelihood-
Contributions in Science, Number 526 Sharkey et al.: Alabagrus of Costa Rica HM 33
Pharpa sp.
19080 © Alabagrus arawak (4)
99 Alabagrus axelhausmanni (1)
82 100 :
98 = too 4 Alabagrus donnai (2)
58
84 gg |A labagrus donlafontainei (5)
1 Alabagrus scottmilleri (3)
95 Posterior prob. x 100
62 190 = Alabagrus cuna (8) Nodal SuppBoer t). (cid:8212)(cid:8212)(cid:8212)ML+ (cid:8212)bo(cid:8212)ot(cid:8212)str(cid:8212)ap(cid:8212) (cid:8212)
400 ~ (cid:8212)(cid:8212)(cid:8212)_] Alabagrus kaydodgeae (3)
86
ie <] Alabagrus cara (8)
Alabagrus bernardoespinozai (1)
400 400 295 20 Alabagrus bobrobbinsi (6)
94 oy (cid:8212)(cid:8212)(cid:8212)__] Alabagrus karensharkeyae (10)
(cid:8212) 100 <=] Alabagrus markmetzi (5)
Bs 90 Alabagrus malcolmscoblei (1)
ino Alabagrus marcepsteini (2)
4100 (cid:8482) Alabagrus iankitchingi (2)
58 a Alabagrus nickgrishini (6)
oo Alabagrus sp. 1 (2)
a 19 (cid:8212)(cid:8212)] Alabagrus nicoya (8)
400 i (cid:8216)| Alabagrus andresfreitasi (4)
* 2 (cid:8212)(cid:8212)(cid:8212)__] Alabagrus andywarreni (5)
mi ; om Alabagrus arua (9)
1] Alabagrus sp2.( 9 )
Alabagrus watsoni DHJPAR0028282
<0 |A labagrus watsoni DHJPAR0028308
| Alabagrus watsoni DHJPAR0028286
4 Alabagrus watsoni DHJPAR0042827
Alabagrus watsoni DHJPAR0023286
101 Alabagrus lindapitkinae (2)
<] Alabagrus watsoni (30)
+ (cid:8212)(cid:8212)] Alabagrus bobpoolei (18)
266 a (cid:8212)(cid:8212)] Alabagrus cocto (13)
92] 93 4a0D0 $$} Alabagrus yuanmaofangi (6)
58 100 =] Alabagrus johnbrowni (29)
1100 (cid:8212)_] Alabagrus mEae culipes (14)
100
99 Alabagrus janzeni (90)
Alabagrus yuchinkengae (1)
99
72 ora Alabagrus brianharrisi (2)
a Alabagrus roibasi (7)
Alabagrus ramyamanjunathae (1)
10? Alabagrus jennyphillipsae (3)
98
2S Alabagrus texanus (10)
89
Alabagrus ixtilton (1)
96] 6g
Alabagrus sp. 3 (1) H806
a (cid:8212) Alabagrus isidrochaconi (19)
100 Alabagrus genemonroei (1)
<(cid:8212)] Alabagrus almasolisae (24)
| Alabagrus sp. 4 (1) H11326
2 {| Alabagrus tanyadapkeae (4)
+00_(cid:8212)(cid:8212)(cid:8212)(cid:8212)] Alabagrus jeanfrancoislandryi (74)
100 Alabagrus paulgoldsteini (1)
4 me {| Alabagrus ekchuah (4)
as 100 __(cid:8212)(cid:8212)} Alabagrus jimmilleri (6)
69 ~~ Alabagrus englishi (3)
a ~ Alabagrus paulthiacourti (2)
70 Alabagrus sanctus (1)
93 Alabagrus masneri (1)
e (cid:8216)é Alabagrus sp. 5 (1) H14623
a <=] Alabagrus imitatus (6)
oT ee (cid:8212)] Alabagrus paulheberti (3)
100 Alabagrus donharveyi (9)
Alabagrus mariaheikkilae (1)
100
ig 96 (cid:8212) Alabagrus kiethwillmotti (23)
100 =] Alabagrus combos (52)
100
72 ass Alabagrus sp. 6 (1) H12099
82 (cid:8216)oe <] Alabagrus jeanmariecardioui (51)
100 os <] Alabagrus jackiemillerae (24)
0.02 substitutions/site 82 16020 Alabagrus fernandodiasi (31)
Figure 1 Tree of highest posterior probability from a 10 million(cid:8212)generation Bayes analysis of CO/. Species are collapsed (when possible) into single terminals (terminal
triangles), with the number of OTUs for each collapsed species in parentheses. The length of the triangles represents the branch length from the node to the tip of the
longest branch.
34 HE Contributions in Science, Number 526 Sharkey et al.: Alabagrus of Costa Rica
based analyses. We conducted a 50-replicate ML search for the tree of RESULTS
highest log-likelihood and a 200-replicate ML bootstrap analysis
PHYLOGENETIC TREES
(Felsenstein, 1985). Both analyses used the default settings.
A Bayesian inference (BI) phylogenetic analysis was also conducted The tree of highest posterior probability from the Bayesian analysis with
species collapsed into single terminals (when possible) is shown in
on the CO/ data set with MrBayes (v. 3.1.2; Huelsenbeck and Ronquist
Figure 1, including nodal support values from the Bayes and ML
2001; Ronquist and Huelsenbeck 2003). As in the ML analyses, the
analyses. We also present four supplemental trees: Bayesian tree of
data were partitioned by codon position. To allow each partition to have
highest posterior probability (Supplemental Figure 1), Bayesian
its own set of parameter estimates, revmat, tratio, statefreq, shape, and
consensus tree (Supplemental Figure 2), tree of highest log-likelihood
pinvar were all unlinked during the analyses. To obtain the most
(Supplemental Figure 3), and ML bootstrap consensus tree (Supple-
accurate branch length estimates possible, the option prset ratepr=vari-
mental Figure 4).
able (which assigns a separate branch length parameter for each
partition) was employed as per the recommendations of Marshall et al.
PHYLOGENETIC CONSIDERATIONS
(2006). A temperature (t) setting of 0.01 was used because there was
The phylogenetic tree in Figure 1 does not have well-supported basal
insufficient chain swapping in a pilot run with the default value t=0.1.
branches. Its primary purpose was as a source of evidence to delineate
With t=0.01, the average chain swapping value was 55%. Two
species. Nonetheless, several lineages are worth noting. Alabagrus
independent, simultaneous BI searches were run for 10 million
fernandodiasi, A. jackiemillerae, and A. jeanmariecadioui form a clade,
generations, saving a tree every 1000 generations, with four search
and all members are similar in color and morphology (BIPP=81,
chains each. The average standard deviation of split frequencies was
MLBS<50). None are represented by males, which is unusual
0.0235 at the completion of 10 million generations. The log-likelihood
considering each is represented by 24 or more specimens. It may
values of the chains in both analyses plateaued just above (cid:8212)10,000 by 3
represent a parthenogenetic clade. Three species, Alabagrus donlafontai-
million generations and did not go any lower by the completion of the
nei, A. cuna, and A. scottmilleri, are distinctly sexually dichromatic, and
run. The likelihood of best state for the (cid:8220)cold(cid:8221) chain of run 1 was
all are found in a clade composed of five species (BIP=84, MLBS<50).
(cid:8212)9911.85 and that for the (cid:8220)cold(cid:8221) chain of run 2 was (cid:8212)9912.76. The
One of the other species, A. axelhausmanni, is represented by a male
5000 post(cid:8212)burn-in trees from each run (10,000 total), using a 50%
only as the holotype, and the other member of the clade, A. donnai, have
burn-in were used to calculate the majority rule consensus tree using males that are darker (orange rather than yellow) than females
PAUP* (v. 4.0810; Swofford, 2003). The data set analyzed herein, (BIPP=99, MLBS=82). Overall it is common to have structural sexual
including the MrBayes command block and Garli configuration files are dimorphism in members of Alabagrus. In particular, sculpture of the
available from the authors upon request. propodeum is often much more pronounced in males than females.
KEY TO COSTA RICAN ALABAGRUS
If you know the species of the host caterpillar it might be best to use the search function on electronic versions of this publication and find the species
of Alabagrus that is (are) on the host, and check your specimen against the images and description. It is doubtful that this key will be very useful
outside of Costa Rica and adjacent countries.
1A. Gena right angled or acute posteroventrally wo... lees eeeeeeeeeeees
1B. Gena rounded or with an obtuse angle posteroventrally .............
Couplet 1
Contributions in Science, Number 526 Sharkey et al.: Alabagrus of Costa Rica Ml 35
ZOWA aW icsoscutuIMmrinostiy -OMentite yainlelalMlGun etteraubierteobteeta cheers etieereouteesmeuuteoupeec whae vtiatea ostantae ohsasabaen siatcesndir setaien omatualit eea aeR t an3
POWb enie soscutumiemiastivaqtsenitttely ap ales, sdanatsSacahstens sees Nest nate Naca sath mes haten deeds nici seal aa Taste Stat asf eee Teese agate ean ated Sac ead 63
Couplet 2
3(2)A x Forewine-almost-entitely infuscate; ignote- small: clear) patches posteriads stie (as i. scacoraeesnst deen svdetecwotde onset decvensene Meine stati Madtcedndate 4
3(2)B. Forewing. patterned iavith large clear, white, (cid:8216)Or-yellow \ateass.....4...-csnnconnsiaecnendenesenedend doneiertedenndentsneetenndenensvisivutearilsiesteeigsddvesiveugaredlostes 18
Couplet 3
4(3)A . Pitsttengum: vatyinie; trom weallytconvex-to witha rounded lomo tii tial. Bulger. ecixn ave devine <ddewseesgtedataigie AglgiadabalgiaestAaladglheMdeMlOeghhe sda ke5
4(3)B e Eiist tereuna swith well-defined median: loneitudinal tear ittay Sees. sss. feleegeustes te etegetussaeness hates ese sebesnlosthsind ssuhvvah iva eabendsvoubesteuebuoiiet 14
Couplet 4
DANa a PrOpOGien RAG AIG uc pasnts imate Bits imate Ruta imate itceleta AMcaleti at lathes A leat led ed ds cdata Mit etal ice atl lela a tledist l tledaA Neda tl bela Al Noda te 6
DICIEIB : «tPOo COUu irte pak es whet hase. ahd th Maser ahd bh aie ah bh Rasen. ahd bias os eal sah need adie ennui a ete A lateo O ate A et Os ea ees lea 8
5(4)C SEr opodernn aiiel aise anteriotly, paler POSter Only -: 220.228 ..c.sasueq! suihnasatetieneatteessaSateresateteieaantseaetSe gaaSttae aAaAdOa MiEtReSa SsNte ,A . combos
Couplet 5
36 M@ Contributions in Science, Number 526 Sharkey et al.: Alabagrus of Costa Rica
6(5)A. Hind femur with weak sparse punctures ventrally, little or no more heavily sculptured than dorsal surface... eee eseeseeeeeeeeeeeees 7
6(5)B. Hind femur heavily sculptured ventrally with aciculations or rugosities, distinctly more heavily sculptured than dorsal surface
hele Aithattt Mecdi tal le lacs acett ect oat be etd Retail sent a octal cana OA lelao ol Shes eA aslalnt M aicand ida ided sarees ada bleicA , donharveyi un. sp.
Couplet 6
TO)A s Media t,t eteite: Ui MAattOWE r. ccnccctecnoneph cenensptcenceap' centnnp cqnchapcertsacotesbberorsahgevssabberotsahgenstespberotiahgerssvchberotvehserssvchsecotpehees A. leedyeri n. sp.
ALGB e Median tergites licwidier sens. sassescsse are staseeresteus anes tans teste eavstecatacaSanayiis aadts seeeeaiss Paslhs seers es als avee Pia esals sveeriiaaees A. johnsharkeyi n. sp.
Couplet 7
SSCS ENA eta Re ee Palle ans ae en a ae Be oR So eng a ae a oe oR Oe RR Se 9
86)B ..M edian: tetvite Sopale anteriorly; melanie posteriory y ok on an,S e oak oe sona ie, os oem stats tee Seed et ee av Te A. watsoni
Couplet 8
9(8)A. Hind femur with weak sparse punctures ventrally, little or no more heavily sculptured than dorsal surface... eee eseseeseeseeeeeeeeees 10
9(8)B. Hind femur heavily sculptured ventrally with aciculations or rugosities, distinctly more heavily sculptured than dorsal surface ......... 1
Couplet 9
Contributions in Science, Number 526 Sharkey et al.: Alabagrus of Costa Rica Ml 37
IO(D)A . Fie coxa embirely: pale acc etanpiat «tdamtesctoestetatiatte ccbtnpluscadethenelanphet aatamhsacasebbeset tape attedincchtcbs nekanase asnnashenohaatheashanbhaaiinasioaohaatsheeun vamnsahstniaon s J
10(9)B.! Hind. coxa pale medially with-2-lareesmelanie patch ror vertical. sirip) laterally. <......ce.c.cnscessncsscssiecsestotesvansssnceaseneesestceaaniss :A . sarapiqui
Couplet 10
kel O-ALt Boca ackecame ellow-. nie cha tues sranereeetas PhatsracbuntstuePeiesteor be eeeuermteaedereatis resid ankeapiaatmnaneie his eased keapuonele aphanthounaatiaapkantyeapmonntannede A. donnai
LbtlO}B. Bodyblack-and "ted te: black. ame Oranges «cain mvaaestaestn assess mara mincaeaanotseen sian ean sian aeaae Menanate. A. jimmilleri n. sp.
Couplet 11
I2OVA- Penultimate metasomal. tereumi (oftempmore terea) mie lane cdr ceca ee ec ade eed hit ded te ot detec dene satin ide Sarin ytvcutsan cteyeiddesersiee A. combos
12(9}B. (cid:8216)Penultimate metasomal'teretin: pale, reddish orange to yellow. <tc cetnces. sce seunserseasvcdebedvsdeubccrsseesesuncdbbatnsddelsabeddoberonddetubiscevbestosblosees(cid:8217) 13
Couplet 12
38 M@ Contributions in Science, Number 526 Sharkey et al.: Alabagrus of Costa Rica
13(12)A. Precoxal sulcus with one or several distinct foveae posteroventrally, with or without a smooth groove extending anteriorly; pale
JeNeg R age) Wl61 785125 4d Ue) eRe Rien re EREr y PERER EPRE PER Sa le Mr rt 4 bt i beta Pe A. donnai
13(12)B. Precoxal sulcus distinct and foveolate ¥2 or more length of mesopleuron; pale parts of body orange........sssseeseeeeseeeeeeees A. arawak
Couplet 13
14(4)A. Hind femur with weak sparse punctures ventrally, little or no more heavily sculptured than dorsal surface .......eieseseseeseeseeeeeeeeees 15
14(4)B. Hind femur heavily sculptured ventrally with aciculations or rugosities, distinctly more heavily sculptured than dorsal surface
SI iene rites cr clam, REL Wo iia beh eed og Pe TCO oR Re RIINS cf: REET 12 ORNL eh EE A, axelhausmanni n. sp.
Couplet 14
PUA oktm C eft UTAUTIO SELVAO E SeTMeSILTY : {PALS Om ae buenH ut vs Biehd uet ts eed ret Heat yeah ar Tah tin aL hee ee ea tT A. bernardoespinozai n. sp.
15014)B . Hind femur mostly os-entitely melanie): s-.sieek, ace Seecnk ehbs ost eeh saas east ost sear p aah eM See L at ee Rea a sha MAU sect aeO MI attend 16
Couplet 15
GGA, Whechan iterate, «Bw Cor tn aial Oe <4. 06. sscantse nics asetajsaadh'e nce hance odds daldasaeeddlesnsleadince snindaslad ages ante haba duane lsaiedeets Taya taateteasanameksnremmts's A. miqa
POCIS)IB : Median terete sito nce tatk aty td ete Nt ete Rate etc te ae ene ed et Hal al tT as AR adie gh Bets ena Be ae ben gf
Couplet 16
Contributions in Science, Number 526 Sharkey et al.: Alabagrus of Costa Rica HM 39
E76;3:1232)A naM ecianisterette(cid:8217) 2p ack a ckieueh wets naa site ok Reaes Sic oh en ak Atha oh Soa Aes koa STR MAR eRe, ST PR A. bobrobbinsi n. sp.
WA33G1, 92 ) Bio Wied iametetet e 2PreOdra tyig e. xcch macs dete sustlesandiz dea usd dese sue Aeeaocsd hestnisb Aste este sineise duttnestdee in aE Mate A. karensharkeyae n. sp.
Couplet 17
L8GyANiPropodeumsareolate, with atleast one clesed mateol a. st .gtheiecdte hoc avket ose abe decd ace duet iiehecah tdctecenscutacey déetudabaccbeceptcstaleeiibecaecctelenttecttsicien 19
[8()B. -Propodeum*lacking-completearedlae, usually mostlyor-completely, smooth)... ....<.-ceesceoneosvotlos ieceiacsehssoceuscdceaessestastesescubtetshesstsoskens s 4]
Couplet 18
TOUS VAS Telecod e ritibely est tnel nate ae oct aca sa raaieeste oe sprec en anata mera vet ohvh te eed doe are te BR pet eR rRayee ie tiie Renate a ne, Seid a Se 20
1901'S)B eli eadewath at least some tracessor paleveOot as... seudeane snewansentsaerdenteane scoters dent vero dentienedeneaepedentseee syes tes aves encntess A. kaydodgeae n. sp.
Couplet 19
20@9)A :Penultimatesmictasoimalyteroutn «(ortinoe)= nicl Ante: wachaconse te cowasptaetete he amdasal RAPA ETT. Sehr AOER Oh hela Ps Gece kOe a eh g. 0h.) 21
20¢19)B;, Penultinvate metasomal teroum.-pale,<reddish= orange toryellowe. <a ccs cecd-wceuese onivesecducste «ihesiiuerusitae i ccavuesesinuemuen A. paulsharkeyi n. sp.
Couplet 20
40 @ Contributions in Science, Number 526 Sharkey et al.: Alabagrus of Costa Rica
21(20)A. Hind femur with weak sparse punctures ventrally, little or no more heavily sculptured than dorsal surface... sees eeeeees 22
21(20)B. Hind femur heavily sculptured ventrally with aciculations or rugosities, distinctly more heavily sculptured than dorsal surface .....38
Couplet 21
22(21)A. Precoxal sulcus with one or several foveae posteroventrally, with or without a smooth groove extending anteriorly... oa)
22(21)B. Precoxaltsulcus; longer, distinct and foveolate sormnoré length of mesopleuron in. <.0...ceessecucceednrcscencetvodeedoonecncancsedchisecdioanncdanettie s 55
Couplet 22
23(22)A. MetapleurotisinostlyOrsertitel ya nie anne. dd sa ncived ie eect ccd one debe vst Su aes Asee se tise hate eae c Ml taen a asta eo ca Ne ta ie ans 24
23(22)B. 2vetapleuron anostlycrrentitclye paler acc aenesane dent cane sereneneten vere denueiene eivinns le Siats lr sive hve ide neat dale St'e te. olase loMat hile oi oboe Shoei us st delestvete dish 30
Couplet 23
2423)P RP ro pOdetm etre Ate tc sce ate obekc ca fds rch hada oth st obuhc otad e ovatU pialinadlda vedo ahead tpn draalesttieanie sles Uarialincdiee- drs derabedl Odea urslbesstadites beg Gerstner ae 25
DAN DS) BME POCCU Ma pal Cartage ze adage da rlagly de gla eda g Lae decd tp cae sthaeos dad las dadbaco nal tlsy tduibaco s tdsthdapeagty de villet ttes éuihaco h dattlss duibac os dal tlss tdusbacoh dey tlss tdustacoh teeta 28
24(23)C . Propodetnr. melamic<atitertorlys< pale sp OstertOLs f <anpscpsanncdss deve dcne esls aclu s token dochS c ooaesk Sate dc oe heatd oce dcge deelS avas detent ede deaes A, almasolisae n. sp.
Couplet 24